flora i,...flora malesiana,series i,volume 13 (1997)1-42 1 rafflesiaceae w.meijerlexington, u.s.a.)1...

Flora Malesiana, Series I, Volume 13 (1997) 1-42

1

Rafflesiaceae

W. Meijer Lexington, U.S.A.) 1

DISTRIBUTION

As defined here the family is subdivided into two subfamilies with a total of nine genera

and about 40 species, mainly in tropical regions, although some species occur in sub-

tropical and even temperate parts of the world. The three genera of the tribe Rafflesieae

of the subfamily Rafflesioideae are only known from Indomalesia, the other two tribes

of that subfamily (Cytineae and Apodantheae) occur in America, Africa, the Mediterra-

nean region (including Iran and Iraq), and SW Australia. The subfamily Mitrastemoi-

deae has only one genus,Mitrastema, originally found in E and SE Asia, but later also

recorded from Mexico, Guatemala, and Colombia(Meijer 1993).

HABITAT

All Malesian species are restricted to moist, evergreen, lowland or montane,primary or

adjacent secondary forests.

1) With contributions by R.W. J.M. van der Ham, Leiden (palynology), and R. Hegnauer,Leiden (phy-

tochemistry). Most original drawings are by Janis Atlee, Lexington, one by Herbert Lee, Tenom

(Sabah).Photographs are from different sources as indicated. Inclusion of the colour plates was made

possible through a substantial financial contribution by the author.

Rafflesiaceae Dumort., Anal. Fam. Pl. (1829) 14; Koord., Bot. Overz. Raffles. Ned.-

Indië (1918) 1-128; Meijer in Kubitzki (ed.), Fam. & Gen. Vasc. Pl. 2 (1993) 557-

563; Hansen in Fl. Thailand2 (1972) 182-184; in Fl. Camb., Laos & Vietnam 14

(1973) 59-64; Kiu & Wing in Fl. Reipubl. Pop. Sin. 24 (1988) 246-248; Dell in

Fl. Austral. 22 (1984) 147-150.

Parasitic plants, rootless, without chlorophyll, always rich in tannins, monoecious or

dioecious.Endophytic body as strings (chains) or plates of cells insidehost plant. Flow-

ering buds or flowering branches vascularized, bursting through the cortex of the host

and terminating in a single flower or in short racemes or spikes, with series of scales

(bracts) at the base of the buds. Flowers uni- or bisexual; perianth single (perigone),

often tubularor saucer-shaped at base, in Rafflesia and Sapria the apex of the tube part-

ly closed by a diaphragm; stamens grouped around a central column representing part

of the pistil in female and bisexual flowers, often joined with that column; ovary uni-

locular, in general inferior but in some cases semi-inferior or superior (Mitrastema),

with 4-6 or numerous parietal placentas. Fruits berry-like. Seeds minute, surrounded

by pulp, testa hard, often thickened and pitted, embryo few-celled, undifferentiated.

Flora Malesiana, Ser. I, Vol. 13 (1997)2

ECOLOGY

The ecology of Rafflesiaceae is of course closely interwoven with the host-parasite rela-

tionship, the chances of survival and the specific ecological requirements of the host

plants, pollination efficiency, seed dispersal and dormancy, and germination chances.

Between the production ofabout 100-150 seeds in a peppercom-sized fruit of Pilostyles

and a fist-size fruit of Rafflesia with several millions of seeds exists an enormous range

of seed production per flower.

Mitrastema is most consistently reported as parasitizing the roots of.Fagaceae: Quer-

cus, Lithocarpus, Trigonobalanus, and Castanopsis and, ifcollectors' notes are accurate,

occasionally adjacent tree roots of other families. All three generaof the tribe

(Rhizanthes,

Rafflesieae

Rafflesia, Sapria) are obligate parasites of the genus Tetrastigma in Vita-

ceae. Where species names are mentioned in the treatment, they must be considered as

preliminary, since identification of herbarium specimens is difficult and nomenclature

uncertain.

Literature: Banziger, H., Nat. Hist. Bull. Siam Soc. 39 (1991) 19-52; ibid. 43 (1995) 337-365.

TAXONOMY

Since Dumortier (1829) most systems of plant familiesof the world have treatedRaffles-

iaceae in a wide sense. Lindley (1836) and Richard (1838) considered Cytinaceae to be

a separate family with Pilostyles as part ofthe Rafflesiaceae and the very closely related

Apodanthus in the Cytinaceae. Van Tieghem (1890) elevated Apodanthaceae to the rank

of family and Makino (1911) put his new genus Mitrastema in its own family. More re-

cent accounts, as Thome (1992), Beaman et al. (1992) and Meijer (1993) treat the fam-

ily in a wide sense. Takhtajan et al. (1985) consideredRafflesiaceae sensu lato as a

heterogeneous group, both macromorphologically and palynologically, and expressed

the idea that the four tribes recognized by Harms (1935) deserve to be treated as separate

families:

Rafflesiaceae: Rafflesia, Rhizanthes,

Apodanthaceae: Apodanthes,

Sapria

Pilostyles

Cytinaceae: Bdallophyton, Cytinus

Mitrastemaceae:Mitrastema.

Studies of ribosomal RNA of parasitic plants carried out by Dr Daniel Nickrent in

Carbondale (University of South Illinois) are still preliminary but already support the

removal of Cytinaceae from the Rafflesiaceae and ultimately may support the family

system of Takhtajan et al. (1985). The ribosomal DNA data (Nickrent & Duff, in press)

make it clear that the tribe Cytineae deserves to become a family at its own. Results on

Apodanthes and Pilostyles are not yet available but it is quite possible also that they can-

not easily be compared with the Rafflesia tribe (or family sensu stricto).

Solms-Laubach (1901: 7) warned not to rush to conclusions about the taxonomy and

relationships ofparasitic plants, which couldhide theirreal ancestry in theirstrong reduc-

tions and adaptations required to survive as parasites. We can safely restrict ourselves

Meijer Rafflesiaceae 3

here to the tribe Rafflesieae and the subfamily Mitrastemoideaeand leave it to future re-

search as to how closely or distantly they might be related.

Literature: Beaman, R.,et al., in: G. Ismael (ed.), Proc. Int. Conf. For. Biol. & Cons. Borneo (1992)

109.— Brown, R„ Ann. Sc. Nat. 2,1 (1834) 369-370; Trans. Linn. Soc. London 19 (1844) 221-239,

5 pi. — Dumortier, B.C. J., Anal. Fam. PI. (1829) 14.— Harms, H., in Engler & Prantl, Nat. Pflanzen-

fam., ed. 2, 16b (1935) 243-281. — Lindley, J., Nat. Syst. Bot., ed. 2 (1836) 389-392. — Makino,

T., Bot. Mag. Tokyo 25 (1911) 254-257, pi. 7. — Meijer, W., Rafflesiaceae, in: K. Kubitzki (ed.),

Families & Genera of Vascular Plants, vol. 2: K. Kubitzki, J.G. Rohwer & V. Bittrich (eds.), Flow.

PL, Dicot. (1993) 557-563. — Nickrent, D.L. & R.J. Duff, Proc. 6th Parasitic Weed Symp. April

1996, Cordoba, Spain (in press). — Richard, A., Nouv. Elem. Bot., ed. 6 (1838) 1518-1520.—

Solms-Laubach, H., in Engler & Prantl, Nat. Pflanzenfam. 3, 1 (1889) 274-282; in Engler, Pflanzenr.

5 (1901) 1-19, 13 illus.— Takhtajan, A.L., N. Meyer, R. & V.N. Kosenko, Bot. J. Leningrad 70

(1985) 153-162. —Thorne, R.F., Aliso 13 (1992) 365-389.—

Van Tieghem, P.E.L., Traite Bot.,

ed. 2 (1890) 1577-1579.

PHYLOGENETIC AFFINITIES

As suggested before, considerations of this subject are still very speculative and depend

on how natural the family in a wide sense really is. Takhtajan (1980) placed the two fam-

ilies Hydnoraceae and Rafflesiaceae in the order Rafflesiales in the superorder Rafflesia-

nae (subclassis Magnoliidae). The closest relationship may well be with Aristolochia-

ceae which already Linnaeus(1753), Robert Brown (1821) and Solms-Laubach (1901)

consideredpossible or likely. Nickrent & Duff (in press) in their preliminary work link

Hydnoraceae with 'paleoherbs' like Aristolochiaceae. The position of Rafflesiales be-

tween Santalales and Celastrales in Cronquist's system (1981) mightbe at variance with

the evidence fromfloral morphology and pollen morphology. As mentionedearlier Takh-

tajan (1985) proposed to raise all the tribes from the system by Harms (1935) to the rank

of family, based on his studies of pollen structure. Thorne (1992), however, stuck to

Takhtajan's family system of 1980 but ranked his Rafflesianae between Nymphaeanae

and Caryophyllanae. The cladistic analysis of Beaman et al. (1992) was also based on

Takhtajan's circumscription (1980) ofthe family and hinges on the selection ofthe most

primitive genus and the most closely related family: Hydnoraceae orAristolochiaceae.

The old family circumscription as followed in this flora can only be maintained when

reversals are assumed.

Literature: Beaman, R., et al., in: G. Ismael (ed.), Proc. Int. Conf. For. Biol. & Cons. Borneo (1992)

109.— Brown, R., Trans. Linn. Soc. London 13 (1821) 201-234, 8 pi. — Cronquist, A., Integr. Sys-

tem Class. Flow. Plants (1981) 703-704. — Harms, H., in Engler & Prantl, Nat. Pflanzenfam., ed. 2,

16b (1935) 243-281.— Linnaeus, C„ Spec. PI. (1753) 442. — Nickrent, D.L. & R.J. Duff, Proc.

6th Parasitic Weed Symp. April 1996, Cordoba, Spain (in press). — Solms-Laubach, H., in Engler,Pflanzenr. 5 (1901) 1-19, 13 illus.

— Takhtajan, A.L., Bot. Rev. 46 (1980) 225-359.— Takhtajan,

A.L., N. Meyer, R. & V.N.Kosenko, Bot. J. Leningrad 70 (1985) 153-162.— Thorne, R.F., Bot.

Rev. 58 (1992) 225-348.

VEGETATIVE MORPHOLOGY AND ANATOMY

The vegetative parts are a kind of tissue mass often compared with fungal mycelia. The

cells have remarkable large nuclei and form the so-called endophytic system within the


stems and roots of the host plants, especially inside the living bark close to the cambium

of the host. From there, special strings (so-called sinkers) can penetrate through rays of

the host into the deeper lying xylem, while these cell strings inside the host cambiumin

general keep pace with the secondary growth ofthe host. In Pilostyles thurberi the endo-

phytic body stays at a uniform distance from the apical meristemof the host (Rutherford

1970), in the Cytineae generally near the ground level and in the Rafflesieae in the roots

as well as high on stems, though in general close to the ground. When the host cam-

bium is reactivated, it may succeed to put a xylem layer on top ofthe parasite tissues. In

general the parasite can migrate through the rays into the new host tissues, as described

by Forstmeier et al. (1983) for Cytinus.

The simple cell strings do not contain vascular tissues; in many genera these occur

mainly in cell cushions from which flowering structures originate, connecting ultimately

with the staminal or ovary structures.

Solms-Laubach (1869) described the situation in Cytinus where the parasite forms a

kind of tissue sleeve between cambiumand xylem of the Cistus host root. In a rhythmic

way, parts of the parasite tissue are covered by xylem layers, apparently and for un-

known reasons turning the cambium activity off from timeto time. The resulting more

mature closed cylinder sleeve of the parasite consists of a cambiumand two surround-

ing layers of tissue: an inner medullary and outer cortical plate. In the latter many irreg-

ular weak vascular bundles occur. Flowering structures originate insideswollenparen-

chyma tissues and cause gall-like growth above connections with the vascular bundles

of the host plant, in general at least 2-3 years after the first infection of the host. The

host tissues are filled with starch at this place except in a few cell layers close to the

parasite tissue. Theseand similar situations were observed by Cartellieri (1926) in Rhi-

zanthes, Brown (1912) in Rafflesia manillana, Haak (1889), Schaar (1898) and Hunzi-

ker (1920) in Rafflesia patma, Endriss (1902) in Pilostyles ingae, Rutherford (1970) in

Pilostyles thurberi and Meijer & Behnke (unpubl.) in Rhizanthes, while the old classical

study by Solms-Laubach (1875) is still of value. If the family is polyphyletic we are

dealing here with remarkable convergences in growth morphology.

Kuijt et al. (1985) discovered sieve elements in the endophytic body of Pilostyles

thurberi, apparently in a discontinuous system which they considered to be a vestigial

cell type. In older studies such elements were in general overlooked.

Weak vascular tissues are reported from the endophytic bodies of species of Pilo-

styles. Floral tissue cushions contain rings of concentric vascular bundles in Rafflesia

and Rhizanthes.

Most of the tissues are parenchymatous and these cells also never contain starch. In

all genera the parenchymatous tissues are rich in tannins.

Stomata are known from scales of Pilostyles and Cytinus and also from Rafflesia and

Rhizanthes (see Cammerloher 1920). Glandularhairs are known from the bracts of Cy-

tinus and also in some species from the perigone. The functionof the so-called ramenta,

the outgrowths inside the flower tubes of species of Rafflesia and from the upper part of

the diaphragm of Sapria is still unknown; their different forms have diagnostic taxonom-

5Meijer Rafflesiaceae

ic value (Winkler 1927). The osmophoric and nectarial structures of the family still need

more comparative anatomical studies.

Literature: Brown, W. H., Philipp. J. Sc., Bot. 7 (1912) 209-224, pi. 12-21.— Cammerloher, H„

Oesterr. Bot. Zeitschr. 69 (1920) 153-164, t. 3. — Cartellieri, E. von, Bot. Archiv 14 (1926) 284- 311,

7 pi. — Endriss, W., Flora, 91, Erg. Band (1902) 209-236, t. 20. — Forstmeier, L., F. Weberling &

H.C. Weber, Beitr. Biol. Pflanzen 58 (1983) 299-312.— Haak, J., Observ. Rafflesias (1889) 14 pp.

— Hunziker, J., PhD Thesis Freiburg (1920). — Kuijt, J., D. Bray & L.R. Olson, Canad. J. Bot. 63

(1985) 1231-1240.— Rutherford, R. J., Aliso 7 (1970) 263-288.

— Schaar, F„ Sitzungsber. K. Akad.

Wiss. Wien, Math.-Naturw. Kl. 107 (1898) 1039-1056.— Solms-Laubach, H., Bot. Zeitung 27 (1869)

185-190; Abh. Naturf. Ges. Halle, 13, 3 (1875) 40 pp., t. 24-27.— Winkler, H., Planta 4 (1927) 1-

97, t. 1.

FLOWER MORPHOLOGY

Only in the genus Bdallophyton do the flowering structures (inflorescences) have a well

developed long axis. In all other genera they are more or less fascicles or very short uni-

flowered shoots, surrounded by a few whorls of scaly dark brown or blackish leaves.

In Rafflesia and Rhizanthes there are in general 3 whorls of 5 scales (bracts), in Pilo-

styles the whorls are 3-6-merous. Apodanthes has a regular structure of 2 outer bracts,

4 calyx-like bracts followed by 4 perigone lobes and an ovary with 4 placentas. Only in

the tribus Cytineae inflorescences are developed like racemes, spikes or umbels, some-

times with bracts and bracteoles.

Flowers are in general unisexual. In Rhizanthes zippelii unisexual flowers occur be-

sides bisexual ones. Bdallophyton oxylepis has bisexual flowers and B. americanum

unisexual ones on different plants; Mitrastema is always bisexual and protandrous.

Perigones can be partly tubularat their bases and the lobes are imbricate (Rafflesia)

or valvate (Rhizanthes). Rafflesia and Sapria possess as unique organ the diaphragma.

Staminal structures always have the thecae interconnectedin a ring like in Hydnoraceae,

though never in phalanges, and are in various ways connected with the central column.

In some cases in Pilostyles they occur in2-4 rings. The detailedanatomy of the anthers

in various genera shows also some variation (see Fig. 1).

Literature: Koorders, S.H., Bot. Overz. Raffles. Ned.-Indie (1918) 206-215.— Meijer, W., in: K.

Kubitzki (ed.), Families & Genera of Vascular Plants, vol. 2: K. Kubitzki, J.G. Rohwer & V. Bittrich

(eds.), Flow. PL, Dicot. (1993) 557-563.

Fig. 1. Comparison of anther structures in different genera, schematic. a. Rafflesia; Sapria;d. Mitrastema.

b. Rhi-

zanthes;

c.

Drawing Janis Atlee.


PALYNOLOGY

(R.W. J.M. van der Ham)

The most detailed account of Rafflesiaceae pollen is that of Takhtajan et al. (1985),

which includes light and electron microscopic dataof 25 species belonging to 7 genera.

Others deal with one or a few species only. Straka (1978), Straka & Friedrich (1984)

and Valdes et al. (1987) provide pollen descriptions with light and scanning electron

micrographs of a few species of the most diverse genus Cytinus.

Pollen of the four tribes, as recognised by Harms (1935), is so different that it can-

not be easily covered in a single description. Pollen grains ofthe

Rhizanthes,

Rafflesieae (Rafflesia,

Sapria) are subspherical to ellipsoidal, 12-21 |im large monads, with one

distal pore or short colpus. The exine consists of a lamellateendexine (2-4 sublayers)

and a loosely attached, sporopollininous perine-like outer layer. Pollen ofRafflesiaarnoldiialso has a homogeneous ectexine. The ornamentationis finely to very coarsely

rugulate-verrucate.

Pollen grains of the Apodantheae (Pilostyles; pollen ofApodanthes and Berlinianche

still unknown) are prolate (P x E = 18-28 x 10-20 gm), tricolpate monads. The colpi

are long and narrow. The exine consists of a homogeneous ectexine and an endexine

that is often lamellatein its outer part. Locally the ectexine may be differentiatedinto tec-

tum, columellateinfratectum and foot layer. Theornamentationis ± psilate.

Pollen grains of the Cytineae (Cytinus, Bdallophyton ) are monads (Cytinus, Bdallo-

phyton), or they are united in tetragonal, rhomboidal or tetrahedral, calymmate or aca-

lymmate tetrads or, sometimes, dyads (Cytinus). The grains are subspherical to ellipsoi-

dal, subiso- to heteropolar, 11-24 gm large, with 2-4 ± equatorial pori or short colpi

(Cytinus), or (2) 3-4 long meridional colpi (Bdallophyton ). The exine consists of a

tectum, a distinct columellateinfratectum and a nexine. The endexine is very thin, and

recognisable only in the apertural areas. Cytinus pollen is sometimes semi- or intectate.

The ornamentation is verrucate, perforate or ± reticulate in Cytinus, and perforate in

Bdallophyton.

Pollen grains of the Mitrastemoneaeare triangular-ellipsoidal, 25 -28 gm large mon-

ads, usually with 3 equatorial(?) pores. The exine mainly consists ofendexine, which is

lamellatenear the apertures. The ectexine is represented by small isolated verrucae. The

ornamentationis psilate to scabrate.

The heterogeneity of Rafflesiaceae pollen led Takhtajan et al. (1985) to the concept

that the family should be split into four natural families: Rafflesiaceae sensu stricto, Apo-

danthaceae, Cytinaceae and Mitrastemaceae(see also p. 2). They did not indicate if these

families form a monophyletic taxon. The single distal (monosulcate, ulcerate) aperture

and thethroughout lamellateendexine ofthe pollen of the Rafflesieae - primitive features

withinthe dicotelydons - suggest that this group belongs to the subclass Magnoliidae,

near the Aristolochiaceae (Takhtajan et al. 1985). With respect to the monophyly ofthe

Rafflesiaceae sensu lato, it is important to determinewhether the (2-) 3- (4-)porate and

-colpate pollen types ofthe other tribes are 'monosulcate-derived' or 'tricolpate-derived'

(Walker & Doyle 1975), and also if the monosulcate/ulcerateaperture type ofthe Raf-

flesieae represents the original, primitive state of the aperture in Magnoliidae pollen.


Harms, H., in Engler & Prantl, Nat. Pflanzenfam., ed. 2,16b (1935) 243 - 281. — Straka,

H., Pollen et Spores 20 (1978) 162-163. — Straka, H. & B. Friedrich, Trop. Subtrop. Pflanzenwelt

51 (1984) 545-546. — Takhtajan, A.L., N.R. Meyer & V.N. Kosenko, Bot. J. Leningrad. 70 (1985)

153-162. —Valdes, B., M.J. Dfez & I. Fernandez, Atlas polinico de Andaluci'a Occidental (1987). —

Walker, J.W. & J.A. Doyle, Ann. Missouri Bot. Gard. 62 (1975) 664-723.

POLLINATION

The foetid smell of the flowers of Rafflesia attracts carrion-flies of the genus Lucilia

(Docters van Leeuwen 1929; Ross in Meijer 1985; Beaman et al. 1988; Banziger 1991).

The smell is produced by fresh flowers, especially during sunny warm periods of the

day. Beaman et al. suggested they observed greenflies in action in Rafflesia at the upper

part of the gullies in the column leading to the anther cavities between rows of hairs

(Meijer 1985). The actual act of pollination is well illustrated by Banziger (1991), see

Fig. 2. Still nobody has fully documented with a video camera what the flies do: laying

eggs, collecting nectar (Haak 1885) or just using the carpet oframentae as a mating ground

(as insects do inside the odorous inflorescences of aroids: Croat, verbal comm.). The flies

carry the pollen on their backs but have not been observed to use it as food. The actual

source area (osmophore) of the bad odour is also still not yet known. Banziger (1991)

suggested from his own observations that the smell originates from the perigone lobes.

Maybe the stomata described by Cammerloher (1920) have a function afterall.

Douglas Warren Stevens noticed during his exploration ofNicaragua a faint smell from

the flowers in Bdallophyton americanum(verbal comm.) and Rutherford (1966)reported

this also from the masses of flowers ofPilostyles thurberi,, both genera which like Cytinus

and Mitrastemon also produce nectar. Honeyeater birds use flowers of Mitrastemon in

New Guinea as nectar source (Beehler 1994).

Banziger (1995: 352-356) gave interesting new data on solidification of the liquid

pollen ofRhizanthes on the back of pollinating insects and reliquefying by profuse stig-

matic fluid and viability over a period of 3 weeks. He also gave details with more recent

names ofthe pollinating carrion flies now classified as Calliphoridae.

Fig. 2. Schematic pictures offlies picking up pollen in a male flower of Rafflesiakerrii (a, b) and strikingit off on the stigma of a female flower (c, d). ss: stigmatic surface. Reproduced with permission from

Bänziger (1991).


References: Banziger, H„ Nat. Hist. Bull. Siam Soc. 39 (1991) 19-52; ibid. 43 (1995) 337-365.

— Beaman, R., et al., Amer. J. Bot. 75 (1988) 1148-1162. — Beehler, B.M., Biotropica 26 (1994)

459-461.— Cammerloher, H., Oesterr. Bot. Zeitschr. 69 (1920) 153-164, t. 3. — Docters van

Leeuwen, W. J., Trop. Natuur 18 (1929) 43-45. — Haak, J., Weekbl. Pharmacie 3 (1885) 19 pp., 2

pi. — Meijer, W., Nat. Geogr. 168 (1985) 136-140.— Rutherford, R.J., PhD Thesis Claremont

Grad. School, USA (1966).

LIFE CYCLE

If we assume that the species of Rafflesia do not vary much in their life cycles we can

reconstruct from data supplied by Teijsmann (1856a, b, 1858), Docters van Leeuwen

(1929), Meijer (1958 and recent unpublished observations) that the total life cycle in that

genus from seed to seed is about 3-4.5 years. New experiments started in Bogor and in

temperate greenhouses in 1991 with seeds of R. gadutensis from West Sumatra, so far

have not produced results. The life cycles of other genera ofthis family in Malesiaare

still incompletely known, except that Banziger (1995) observed for buds of Rhizanthes

lowii (Rh. zippelii sensu Banziger) that it took an estimated 200-255 days from the

point of breaking through the host tissue at a circumference of3.7 to 4.1 cm to opening

of the flower.

References: Banziger, H., Nat. Hist. Bull. Siam Soc. 43 (1995) 337-365.—

Docters van Leeuwen,

W. J., Trop. Natuur 18 (1929) 43-45. — Meijer, W., Ann. Bogor. 3 (1958) 33-44. — Teijsmann, J.E.,

Nat. Tijds. Ned. Indie 12 (1856a) 279-281; Hook. J. Bot. & Kew Gard. Misc. 8 (1856b) 371-374;

Ann. Hortic. Bot., Fl. Jard. Roy. Pays-Bas (1858) 27-30.

FRUITS AND EMBRYOLOGY

Field observations in 1981 and 1983in the Ulu Gadut area near Padang, West Sumatra

have shown that ripening of fruits of Rafflesia gadutensis takes about 8 months. Two

fmits were monitored for periods of 6-7 months and the eldest had ripe seeds in Febru-

ary 1984. Meijer's guide Satar, who was monitoring this fruitbelow his hut, mentioned

that it was visited by a 'tupai kuning' (most likely a tree shrew), the same species which

was eating his young chickens. In Sumatra as well as Borneo we noticed male buds be-

ing opened by squirrels or/and tree shrews and it looks as if one shrew sitting on the

exit of such a vandalized bud was captured on the lens on Kinabalu near Poring; see

Attenborough (1995). It sits on a bud not at the entrance of a flower as the legend in the

book asserts. The holes made are too small for most squirrels.

From a very obscure report by R.H.C.C. Scheffer in a letter to Solms-Laubach [cited

in Solms-Laubach (1875: 27)] it might appear thatactual pollination is not necessary for

fruit set. Scheffer wrote to Solms (translated): "Only one of the buds made it into a flow-

er, only 3 years after the inoculation. Notwithstanding this was a female one, and with-

out having a male flower nearby it still developed seeds. These were used to infect an-

other plant of Cissus scariosa [= Tetrastigma leucostaphylum] and this was successful."

Solms asserts that Scheffer suspected that the rudimentary anthers might have pro-

duced some pollen. That is very unlikely. Meijer's unpublished field observations

showed that unfertilized female flowers, several months after flowering, had produced

no seeds with embryos. Ovaries with very little pollination may be stimulated to grow


into fruits, but they can be full of seeds without embryos as observed from a fruit of

Rafflesia keithii outside the Kinabalu National Park, August 1993 (seeds tested by

F. Bouman, Amsterdam). Possibly there were other flowers of Rafflesia open in the

garden or on Mount Salak about 120 years ago, to account for Scheffer's observations.

Real parthenocarpy of Rafflesia has never beenproven.

Ovule and seed development stages have been studied for Rafflesia and Rhizanthes

(Solms-Laubach 1898; Ernst & Schmid 1913). Pollen tetrads form according to the suc-

cedaneous system; the tapetum of anthers consists of 2 or 3 layers of cells. The stigma

is in Rafflesia and Rhizanthes situated in a ring around the rim of the column disk, often

more or less papillose; the ovary shows in these genera a maze of placental plates which

fill the ovary cavity. For the development of the embryosac see Ernst & Schmid (1913)

and Olah (1960). The outer integument is reduced. Ovaries grow into berry-like fruits,

with whitish pulpa around the seeds. The seeds (Fig. 3) are small and with hard scales

[see Harms (1935: 255) and Bouman & Meijer (1994)]. Rafflesia patma has a 2-cell

suspensor and an embryo consisting of 3-5 layers of 2-4 cells, often only 6 cells in

total. Endosperm is nuclearand consists of 30-40 cells in R. patma. Endosperm and em-

bryo mostly contain oil, see also Schuerhoff (1926: 526). Mitrastema also has cellular

endosperm, but ovules with only one integument.

Attenborough, D., The private life of Plants (1995). — Bouman, F. & W. Meijer, PI.

Syst. Evol. 193 (1994) 187-212.— Ernst, A. & E. Schmid, Ann. Jard. Bot. Buitenzorg II, 12 (1913)

1-58, t. 1-8.— Harms, H., in Engler & Prantl, Nat. Pflanzenfam., ed. 2, 16b (1935) 243-281.

—

Olah, L.V., Bull. Torrey Bot. Club 87 (1960) 406 - 416. — Schuerhoff, P.N., Zytol. Blutenpfl. (1926).

— Solms-Laubach, H., Abh. Naturf. Ges. Halle 13, 3 (1875) 40 pp., t. 24-27; Ann. Jard. Bot. Buiten-

zorg 9 (1891) 184-246,28 pi.; ibid., Suppl. 2 (1898) 11.

Fig. 3. Mature seeds of (Blume)

Spach (right). Scanning electron micrographs F. Bouman, courtesy Hugo de Vries-Laboratorium, Uni-

versity of Amsterdam.

Rafflesia rochussenii Teijsm. & Binn. (left) and Rhizanthes zippelii


DISPERSAL

Watanabe (1933, 1936) assumed long range seed dispersal ofMitrastema by birds. Ob-

servations of jungle walkers working for Meijer (1958, 1983) at the sites of Rafflesia in

West Sumatra showed that ground squirrels and tree shrews like to eat the contents of

the whitepulpy ripe fruits of these plants. These observations have been confirmed now

by Emmons, Jamili Nais and Ali Briun(1991) and documentedwith colour photographs

froma hide erected in August 1989 near a fruitof Rafflesia keithii on MtKinabalu. The

fruits ofRhizanthes also have white pulp, but they are hidden under the old perigones

and predation has never been observed yet.

In analogy with other root and stem parasitic plants it is assumed that only distur-

bance and damage to the organs of the host can deliver the signal for seeds in contact

with themto germinate (Bouman & Meijer 1994).

References: Bouman, F. & W. Meijer, PI. Syst. Evol. 193 (1994) 187-212. — Emmons, L., et al.,

Biotropica 23 (1991) 197-199. — Meijer, W., Ann. Bogor. 3 (1958) 33-44, fig.; Essays on Rafflesia-

ceae (1983) 1-20.—Watanabe, K„ Bot. Mag. Tokyo 47 (1933)798-805; J. Jap. Bot. 12 (1936) 603-618.

PHYTOCHEMISTRY

(R. Hegnauer)

This mainly tropical family ofparasitic plants is badly known at present from a chemical

point of view (see Hegnauer 1973, 1990). Cytinus hypocistis yielded isoterchebin, an

ellagitannin, as its yellow pigment and pelargonidin 3-galactoside and petunidin 3-gluco-

side as red flower pigments. The sole crystalline compound isolated from acetone ex-

tracts of Psilostyles thurberi turned out to be sucrose. Pollinationbiology of Rafflesia-

ceae is still poorly known. Recent observations with Rafflesia pricei suggest that optical

and olfactory mimicry may offer the clue. Flower pigments and odorous principles em-

anatedfrom flowers attract carrion-flies; they seem not to be compensated for theirpolli-

nation activities by nurture or suitable breeding sites. Most authors are convinced that

Rafflesiaceae belong to or are affiliatedwith Polycarpicae sensu Wettstein. However,

synthesis and accumulationof ellagitannins do not favour such an assumption, because

polymeric proanthocyanidins (i.e. 'condensed tannins') are the characteristic tannin-like

metabolites of Polycarpicae. In this respect Rafflesiaceae are similar to Nymphaeaceae

s.str. which produce ellagitannins and possibly Cabombaceae which produce gallic

acid and probably gallotannins. Therefore it is noteworthy that Takhtajan (1980) has

Aristolochiales, Rafflesiales, Nymphaeales (Cabombaceae, Nymphaeaceae s.str. and

Ceratophyllaceae) and Nelumbonales as numbers 5-8 in his subclass Magnoliidae, and

that in Thome's (1992) classificationAristolochiaceae are incorporated in Magnolianae-

Magnoliales-Magnoliidae and the superorders Nymphaeanae and Rafflesianae immedi-

ately follow Magnolianae. Nelumbonalesand Ceratophyllales are treated by Thome as

orders ofMagnolianae. Synthesis and accumulation of gallo- and ellagitannins may have

evolved in a numberof taxa now considered as outgroups of true polycarps.

References: Hegnauer, R., Chemotaxonomie der Pflanzen 6 (1973)9; 9 (1990) 314-315. Takhta-

jan, A.L., Outline ofthe classification of floweringplants (Magnoliophyta), Bot. Rev. 46 (1980) 225-

359. Thome, R.F., Classification and geographyof floweringplants, Bot. Rev. 58 (1992) 225-348.


KEY TO THE GENERA

1 a. Ovary superior, staminal structure a tube surrounding the pistil with a series of an-

thers above each other. Host plants in general Fagaceae ....

Mitrastema (p. 11)

b. Ovary inferioror half-inferior, staminal structures different.Host plants species of

Tetrastigma (Vitaceae) 2

2a. Mature flower buds oblong. Perigone without diaphragm, lobes 16-18, valvate,

ending in bayonets (elongate stiff pointed appendages), which are hidden in a cav-

ity at apex of the column while flowers are in the bud stage . Rhizanthes (p. 37)

b. Maturebuds more or less globular. Perigone with a diaphragm, the lobes imbricate,

5-10 3

3a. Perigone lobes 5; ramenta only inside the flower tube Rafflesia (p. 13)

b. Perigone lobes 10; ramenta on top ofthe diaphragm . .. Sapria (Continental Asia)

MITRASTEMA

Mitrastema Makino, Bot. Mag. Tokyo 23 (1909) 326 ('Mitrastemma’). —Mitrastemon Makino, Bot.

Mag. Tokyo 25 (1911) 225, orth. mut., nom. inval.; Jochems, Rec. Trav. Bot. Neerl. 259 (1928)

203; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 16b (1935) 274; Yamamoto, Bot. Mag.

Tokyo 50 (1936) 539; Watanabe, J. Jap. Bot. 13 (1937) 154; Matuda, Bull. Torrey Bot. Club

74 (1947) 133; Meijer in Kubitzki (ed.), Fam. & Gen. Vase. PI. 2 (1993) 560; Meijer & Veld-

kamp, Blumea 38 (1993) 221-229.— Type species: Mitrastema yamamotoi Makino (‘Mitra-

stemma’).

Chlorophyll-less endoparasites with fungus-like endophytic body. Stems unbranched.

Leaves scale-like, decussate. Flowers solitary, terminal, bisexual. Perianthmuch reduc-

ed, collar-shaped. Stamens numerous, completely connate into a tube, below the sterile

apical part with several vertical series of rings of c. 10 anthers each; pollen dicolpate,

with reduced ectexine. Ovary superior, 1-locular, ovules numerous, anatropous, uni-

tegmic. Fruit a slightly woody, berry-like capsule, horizontally dehiscing. Seeds sticky,

testa hard.— Fig. 4,5.

Distribution — Two species, one in Asia, from Japan to Taiwan, Yunnan, Bhutan,

Assam, Indo-China, and Malesia: Borneo, Sumatra and New Guinea; a second, very

closely related species (?) from Chiapas (Mexico) to Alta Vera Paz (Guatemala) and

Antioquia (W Colombia).

Habitat— In montane oak-chestnut forests, altitude 1400-2000m.

Ecology — The Asian species parasitic on roots of Fagaceae, probably the American

species too. Pollination by insects and birds, seed dispersal most likely by birds and

small mammals.

Notes— 1. According to Matsuura, J. Fac. Sc. Hokkaido Imp. Univ. V, Bot. 3 (1935)

189, and Watanabe, J. Jap. Bot. 12 (1936) 769, f. 26P, the chromosome number is

n = 20.

2. About the orthography of the genus name, see Meijer & Veldkamp, I.e.


Mitrastema yamamotoi Makino

Mitrastema yamamotoi Makino, Bot. Mag. Tokyo 23 (1909) 326, fig.; P. Royen, Nova Guinea, Bot.

14 (1963) 243-245, pi. 17; Hansen, Bot. Tidsskr. 67 (1972) 149; in Fl. Camb., Laos & Vietnam

64 (1973) 62, t. 9; Tang Shui Liu & Ming You Lai in Fl. Taiwan 2 (1976) 582, t. 414; Meijer &

Veldkamp, Blumea 38 (1993) 227. — Syntypes: Tashiro s. n.,Bando s. n., Yamamoto s. n. (TI holo),

Japan, Kyushu.

Mitrastemon sumatranum Nakai, Icon. PI. As. Or. 4 (1941) 338, t. 113 (‘sumatranus’).— Type; Jochems

s. n. (TI holo), Sumatra, Karo Mts.

For more complete synonymy and references, see Meijer & Veldkamp, 1.c.

Plants about 2.5-15 cm long and up to 3 cm in diameter.Basal part a corky pustular

cupule, up to c. 2 cm high and in diameter, the margin an irregularly low lobed rim;

stems 5 -15 mm diam. Leaves (also calledbracts or scales) in 3-4(-7) decussate whorls,

increasing in length from the base of the plant upwards, broadly to narrowly ovate, at

apex blunt-rounded, (4—)10—20(—28) by (5—)10—15(—25) mm, cream (turning ebony)

coloured with brown spots, the lowest often purplish brown. Flowers up to 20 mm

long, with a foetid smell. Perigone 5-10 by 6-17 mm, truncate or slightly 4-lobed. Sta-

minal tube tubular, wide and globose in the basal part, 14-20mm long, the minute an-

thers in a 2-5 mm broad ring in the upper part, the whole collar early caducous. Ovary

ellipsoid, c. 8 by 3 mm, style ovoid-ellipsoid, 2-8 by 3-6 mm, stigma 2-5 by 4-6 mm;

8-15 radial placentas. Fruit up to c. 20 by 10 mm, opening along a horizontal ring about

halfway. Seeds c. 0.25 mm diameter, with a small funicle, yellowish or dark brown. —

Fig. 4, 5.

Distribution — India, Thailand, Cambodia,Vietnam, China (Yunnan), Japan, Taiwan;

Malesia: Sumatra (Aceh, East Coast), Borneo (Sabah: Mt Kinabalu; Sarawak: 4th Divi-

sion), Papua New Guinea (W and E Highlands, Morobe, Milne Bay Provinces).

Fig. 4. Makino. Full open flower, habit and length section, with indication of

flower parts. True size up to 15 cm high. Drawing by Janis Atlee after the type description by Makino.

Mitrastema yamamotoi


Habitat — Montane oak forests, at altitudes of 1400 to 2000 m.

Ecology — In MalesiaMitrastemayamamotoi is in general reported as a root parasite

of Fagaceae: Castanopsis, Lithocarpus, Quercus, Trigonobalanus [Akuzawa, Paras. PI.

Magaz. 13/4 (1985) 23], but also on species of other families. Flowering is reported on

Mt Kinabalu around April, the fruits ripening in about 4 months [Akuzawa, Paras. PI.

Magaz. 15/1 (1986) 26],

RAFFLESIA

Rafflesia R. Br., Trans. Linn. Soc. 13 (June 1821) 201; Jack, Appendix Descr. Mai. PI. 1 (July 1821);

Jack ex Hook., Comp. Bot. Mag. 1 (1835) 259, pi. 14; R. Br., Trans. Linn. Soc. 19 (1844) 221;

Solms-Laubach in Engl. & Prantl, Nat. Pflanzenfam. 3, 1 (1889) 279; in Engl., Pflanzenr. 5 (1901)

8; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 16b (1935) 259; Meijer, Blumea 30 (1984)

209; in Kubitzki (ed.), Fam. & Gen. Vase. PL 2 (1993) 561. — Type species: Rafflesia arnoldii

R. Br.

Parasitic plants, endophytic body growing like a thallus inside the woody stems and

roots of species of Tetrastigma (Vitaceae). Flower buds sessile, first protruding as a

corky swelling with hexagonal patches, the corky base remaining as a cup-shaped body

(cupule) below the scales on the flowering sessile shoot. Scales (bracts) in a series of 3

whorls of 5 scales, imbricate, at first appearance white, but turning black or dark brown

Fig. 5. Makino. Dried flower on host. Sumatra, Berastagi, coll. by A. Stomps.Photo by F.D. Boesewinkel, Amsterdam.

Mitrastema yamamotoi


after exposure, with prominent veins, inner gradually larger. Flowers (see Fig. 6) uni-

sexual, female ones with rudiments of anthers. Perigone lobes 5, imbricate, reddish,

often with white warts, inserted around an annular, horizontal, pentagonal, central dia-

phragm which has a more or less round opening (orifice) in the centre (see Fig. 7, 8).

Meijer. a. Flower with the small aperture in the diaphragmclearly show-

ing; b. female flower, halved lengthwise, showing central column (ap: aperture in diaphragm; coll:

collum, neck of column; cup: cupula, perigone tube; ov: ovary; proc: processi on apex of disc; ram:

ramenta on inside of cupula and diaphragm; sul: sulcus under disc); c. side view of the column of a

female flower showing outer and inner annulus (ae: annulus exterior, ai: annulus interior); d. section

through male flowers, anther in longitudinal section and seen from lower side of the overhang of the

‘corona’ of the disc towards the sulcus; e. details of ramenta, often branched and with swollen apices.

Drawings by Janis Atlee (a, b, d, e) and Herbert Lee (c), made from colour slides by W. J. de Wilde and

drawings from BO.

Fig. 6. Rafflesia micropylora


Base of perigone tube and adjacent zone of diaphragm (or the entire diaphragm) covered

with variably shaped ramenta inside (see Fig. 9). Central column at apex widened into a

disk, often with processes, below this narrowed into the neck and at the base widened

and surrounded by the 'annulus interior' above a sulcus, the base of the flower tube of-

ten thickened into an 'annulus exterior' which can be almost or totally obsolete. Anthers

with one apical pore, sessile, situated around the overhanging rim of the column. Ovary

1-locular with many placentas. Fruit berry-like. Seeds thick-walled, c. 1 mm long and

with a shorter appendage. — Fig. 6-9, Plates 1-4.

Distribution — About 13 species from the Kra Isthmus of Thailand through West

Malesia, including Luzon and Mindanao in the Philippines. Possibly formerly also in

Bali.

Habitat — In primary and secondary forest, the host Tetrastigma species prefer to

grow in rich alluvial or limestone derived soils. As a result they never occur in areas

with Gleichenia thickets. Moreover they are restricted to altitudes below about 2000 m.

The highest reported Rafflesia is the site on Mt Leuser where Van Steenis found R.

rochussenii at 1800 m. Most of the known localities are around 1000 m or lower. That

may be the reason Rafflesia is now very rare on the rather denuded lower mountain

slopes of West Java.

Some of the best Rafflesia and Rhizanthes sites are foundoutside old primary forests

since Tetrastigma species germinate best in open disturbed soils from the seeds dispers-

ed by birds or small mammals. How well the parasites can survive in severely cut back

climbers is still unknown, but this can now be checked at some sites where silvicultural

operations have damaged populations.

Ecology — Always parasitic on the trailing stems and /or roots of some species of

Tetrastigma (Vitaceae).

Fig. 7. Rafflesia spp. Semidiagrammatic representation of the diaphragms of: a. Rafflesia hasseltii

(7 concentric

rings of very small white warts), and c.

(4 concentric rings, not ofwarts but at same level as surrounding tissue), b. Rafflesia kerrii

(5 concentric rings of white warts). Drawing

by Janis Atlee.

Rafflesia keithii


n: Drawn by Janis Atlee, from different sources in archives W. Meijer (MO).R. arnoldii.

g: R. keithii,R. patma, R.

schadenbergiana,

m:l:R. micropylora,k:R. tuan-mudae,j:i:R. gadutensis,h.R. hasseltii,

e:c: R. pricei,d:R. tengku-adlinii,R. rochussenii,b: R. kerrii,f:R. cantleyi,R. manillana,

Rafflesia. Fully open flowers. For each species a flower of average size is pictured from above,

a:

Fig. 8.


Taxonomy — Winkler [Planta 4 (1927) 1-97] suggested a subdivision based on the

structure of ramenta, which appear to have great diagnostic value in tracing the possible

phylogeny of species in this genus. Comparative research of the ribosomal RNA of sur-

viving species can test this assumption.

Historical review — The first botanical explorer who foundRafflesia in Malesia was

the prominent but ill-fated French surgeon-naturalist Louis Auguste Deschamps. He

sailed from 1791 to 1794 during the great upheavals of the French revolution with the

expedition of the 'La Recherche' in search of the lost research ship 'La Perouse' (Van

Steenis-Kruseman 1950; Van Steenis et al. 1954). In March 1794 the ships, after giving

up the search, were in the harbour of Surabaya in East Java and the commander of the

expedition refused to proceed in view of the situation in France. Deschamps and his

companions let themselves be interned by the Dutch colonial authorities. Deschamps,

who apparently showed great interest in the local language, people, flora and fauna was

Ramenta from inside of perigone tube from 13 species, drawn on scale, a:Fig. 9. Rafflesia. R. patma,

R. keithii, f: R. gadutensis, R. micropylora,c: R. lengku-adlinii, d:b: R. manillana, R. pricei, e: g:

h: R. kerrii, i: R. arnoldii, j: R. rochussenii, k: R. schadenbergiana, l: R. cantleyi, m: R. hasseltii.

Drawn by Janis Atlee, from different photographs in archives W. Meijer (MO).


invited by Governor Van Overstraten to make natural history studies all over Java for

three years, making with the help of special assistants many notes, drawings and collec-

tions and even starting on a manuscript entitled 'Materials towards a Flora of Java'. That

still marvelous islandmust during that timehave been for him a botanical paradise. In

1798 he could return with his treasures to France but meanwhile the British were at war

with the French and blockaded France from the sea. The ship on which Deschamps sail-

ed was taken by the British and all the specimens and notes of Deschamps were confis-

cated.

Years later the manuscripts and some specimens turned up at a public sale at the

Indian House inLondon and were bought by John Reeves, who donated them in 1861

to the British Museum (Natural History). The specimens were never seen again (Van

Steenis-Kruseman 1950). It lasted until about 1954 when the manuscripts could be

studied by C.A. Backer, Mrs and Prof, van Steenis, who to their surprise discovered

that among these notes were sketches of buds and host plants of Rafflesia, most likely

seen by Deschamps on Java, long before the well-known so-called first discovery of

what somebody called that stupendous plant (Mabberley) by Arnold and Raffles in the

southern part of Bengkulu (Benkoelen, Bencoolen) in Sumatra in 1818. There are sus-

picions by Mabberley and Van Steenis (unpubl. letters) that British botanists possibly

knew that Deschamps had found a very strange plant and that therewas a kind of secret

competition going on to see who would first unravel further this wonder of nature.

That might have been one of the reasons why the young and eager botanical explorer-

surgeon-naturalist William Jack, replacing Arnold who died of malariasoon after he did

his discovery, rushed at the instructionsof Raffles a description of Rafflesia as R. titan

Jack, already forwarded to Marsden by Raffles in April 1820, intoprint in August 1820

[see W.J. Hooker (1835) 135, 136] in an Appendix of his Malayan Miscellanies at the

Missionary Press in Bengkulu. Actual publication of this Appendix was kept back while

the Linnean Society took long to print the paper on R. arnoldii R.Br., read by Brown

on 30 June 1820 at the meeting of this prestigious society. Whoever got the name

first published it should be a British botanist. (See also page 35 under the Incompletely

known species.) Rivalry between the British, French and Dutch was great at that time.

The supposition of Merrill (1952) that the names R. arnoldii and R. titan were based

on the same types is wrong. Jack collectedplenty of extra materials for Brown and him-

self after Raffles had forwarded letters and specimens to Sir Joseph Banks, who dele-

gated the description to Brown (Mabberley 1985) and he also established for sure that

the plant was a parasite on a kind of grapevine, later calledthe genus Tetrastigma. Soon

after Jack had assured Robert Brown that his publication was only a kind of backup ac-

tion he died of a lung disease and only a few copies of his Appendix had been sent by

him to Wallich in Calcutta and to Brown, while the main shipment went into flames at

sea not far from the harbour of Bengkulu with all of Raffles's belongings on his way

back to England. Raffles managed to get ashore again and started a recollecting drive

(Mabberley 1985). See also details on Jack in Hook. Comp. Bot. Mag. 1 (1835) 121—

147. He was a marvelous very promising botanist like Arnold and later Kuhl and Van

Hasselt who were less lucky than Deschamps to get out of malaria-infested Java alive.


Just a few years after Brown received letters from Jack about the fruits of Rafflesia

and its host specificity another Rafflesia event unfoldedin Java where since 1818 the

very young and ambitious (only 22 years old in 1818) German-Dutch surgeon-naturalist

C.L. Blume was stationed as inspector ofvaccines and in 1822 as Director of the Buiten-

zorg (now Bogor) Botanical Garden. He used the fortuneof his rich first wife to launch

large expeditions in the wilds of Java, during one of which he foundout about a large

bud and some kind of flower in the Nusa Kambangan Peninsulaalong the southcoastof

Java. He succeeded to find it, first did not recognize it as a flowering plant, but soon

compared it with the excellent illustrations of the Bauerbrothers who worked for Robert

Brown. By the timehe had returned to Hollandthe German plant collector A. Zippelius

had found on Mount Salak near Bogor what appeared to be a second genus related to

Rafflesia. Blume described his new Rafflesia patma and also the new genus and species

which he named Brugmansia zippelii, unfortunately using a genus name already occu-

pied in Solanaceae. In beauty and interesting flowerbiology this genus, now known as

Rhizanthes, equals Rafflesia and the large plates and long descriptions in Flora Javae by

Blumeare very impressive although difficult to read in his elaborate Latin prose (Backer

1921).

Subsequently other discoveries were made: the very small Rafflesia manillanain Lu-

zon (Teschemacher 1841); in 1850 a rather small novel Rafflesia species with a com-

pletely unarmed disk of the central column, collected by an unknown plant collector of

the Bogor garden on the Gedeh-Pangerango outlayers near Cibodas and described by

Teijsmann & Binnendijk (1850) as R. rochussenii. In 1884Rafflesia tuan-mudae was

found by O. Beccari in Sarawak. The Central Sumatra expedition of D.D. Veth in 1884

had the very distinct R. hasseltii as one of its main trophies. It was recently rediscover-

ed by the French botanist-cartographer Yves Laumonierand by Harry Wiriadinatafrom

Bogor.

Solms-Laubach (1891) commented on the two species known from the Philippines

and he described in 1910R. cantleyi from Malaya.

Meanwhilethe few known localities ofRafflesias within easy access from Bogor had

been mined out so much for all sorts of anatomical-morphological research and attempts

to transplant the parasites wholesale attached to the hosts to the Bogor Botanic Garden,

that the original localitiesbecame depleted.

Koorders (1918) described a number of new species from collections by Witkamp

and others; Meijer (1984) added five and Mat Salleh & Latiff (1989) one. In 1992 R.

kerrii was found back in the Malay Peninsula(Wong 1993).

References: Backer, C.A., Trop. Natuur 10 (1921) 124-127.— Koorders, S.H., Bot. Overzicht

Rafflesiaceae Ned.-Indie (1918) 128 pp. — Mabberley, D. J., Jupiter Botanicus. Robert Brown of the

British Museum (1985) 219-238. — Mat Salleh, K. & A. Latiff, Blumea 38 (1989) 111-116. —

Meijer, W., Blumea 30 (1984) 209-215. — Solms-Laubach, H., Ann. Jard. Bot. Buitenzorg 9 (1891)

184-246, 28 pi. — Steenis, C.G.G.J, van, M.J. van Steenis-Kruseman & C.A. Backer, Bull. Brit.

Mus. (Nat. Hist.) I, n. 2 (1954) 51-68, pi. 13.— Steenis-Kruseman, M.J. van, Cyclopaedia of col-

lectors. Flora Malesiana I, 1 (1950). — Teijsmann, J.E. & S. Binnendijk, Nat. Tijds. Ned. Indie 1

(1850) 425-430. — Teschemacher, J.E., Boston J. Nat. Hist. 4 (1841) 63.— Wong, M., Nature

Malays. 17(1993) 124.


Ex situ cultivation—

There are reports by Teijsmann (1850) and J.J. Smith (1929)

about transplantations of Tetrastigma with Rafflesia from their natural sites to the Bogor

Garden. In 1850 H.H.Loudon from Nusa Kambangan assisted with the transfer of

Rafflesia patma [see a letter from Teijsmann to De Vriese, dated 23 July 1852 in De

Vriese (1853)]. This plant flowered from 23 until 25 March 1850 and again the 6th of

October. Docters van Leeuwen (1929) cited the same flowering dates and this proofs

that in fact R. patma was not grown from seeds as he asserted. Rafflesia rochussenii

was added to the garden collection from transplants after 29 July 1850 (Teijsmann &

Binnendijk 1850). By 1856Rafflesia arnoldii had been added to it (see Teijsmann 1856a;

Teijsmann & Binnendijk 1866).

According to a short note in Botanische Zeitung of 15 May 1857 (p. 328), the Dutch

newspaperHaarlemsche Courant of 7 April 1857 stated (translated):"

Rafflesia arnoldii

flowered on 9 February 1857 in the Botanic Garden of Buitenzorg, Java. This flower

came from a plant which grew as the result of Teijsmann's attempts to inoculate a wild

grape vine. The flower was immediately illustrated at the spot." Most likely this is the

one figured by Miquel (1863).

As mentionedabove, Teijsmann (1856a, b) reported in several papers abouthis inoc-

ulation experiment, also in a French version (Teijsmann 1858). Binnendijk, in a letter to

Motley (Motley 1857), claimed that he in fact did the actual inoculationwhich took place

in November 1854. The first buds were visible in August 1855 and as we saw the first

flower appeared February 1857.

Seeds were said to have been taken from a fruit grown in the garden but it is rather

obscure how the flower which produced that fruit was pollinated. Also the subsequent

history as reported by Solms (1875) and Beccari (1875), suggesting that this plant was

bisexual, self-pollinating or parthenocarpic is very confusing as already explained above.

It shows that careful documentationand labelling in situ and good record keeping is ab-

solutely needed in such experiments.

In 1866, there were three species of Rafflesia (R. patma, R. arnoldii and R. rochus-

senii) at the Bogor Garden according to the Garden Catalogue (Teijsmann & Binnendijk

1866). Docters van Leeuwen (1929) citedold mantri Noerkas as remembering a flower

in 1879.

The situation in 1883/84 was describedby Solms (1891). Beccari (1875) reported to

have seen flowers of R. patma in Bogor when he visited the Gardens between 30 Janu-

ary and 2 February 1872.

Beccari saw R. arnoldii in the Bogor Garden in 1874 (Beccari 1875) and reported that

the species flowered every year. Reports from Docters van Leeuwen can be projected

back in time around 1889 and around 1898 according to J.J. Smith (1929). Raciborski

made according to Smith new attempts to transplant R. patma from Nusa Kambangan.

The last year R. rochussenii and R. patma flowered in Bogor was 1929 (Docters van

Leeuwen 1929; J.J. Smith 1929). Since 1929 no furtherattempts were madeby anybody

connected with the garden since localitieswere depleted, partly due to over-collecting or

totally destroyed by land cultivation or protected in tiny nature reserves (see also Brewer

1918).


Docters van Leeuwen wrote in a Dutch article (1929) that his horticulturistP. Dakkus

repeated a seed inoculation experiment in August 1924with seeds of Rafflesia rochus-

senii from a fruit supplied by Ader from Mt Garut. This was done in the Fern-Bromeliad

Garden on stems of Tetrastigma lanceolarium (now T. leucostaphylum). In January of

1929, 4.5 years later, a flowerappeared as a result of this. Dr. Frits Went made photo-

graphs of it.

Rafflesia rochussenii seems to be the only species of the genus that has ever flower-

ed in a greenhouse in the temperate zone. De Vriese (1853) illustrated this species from

a plant grown in the Leiden Botanical Garden. The infected host plant must have been

transported in a wardian case by ship all the way from Java. I have not been able to find

any special report about the actual flowering date in a local newspaperof that time nor a

reference in Botanische Zeitung.

References: Beccari, O., Nuovo Giorn. Bot. Ital. 7 (1875) 70-75. — Brewer, F.W. J., Meded. Natuur-

mon. Ned.-Indie nr. 2 (1918) 19-21.—

Docters vanLeeuwen, W. J., Trop. Natuur 18 (1929) 43-45.

— Miquel, F. A.W., Choix des plantesrares ou nouvelles cultivees et dessin6es dans le Jardin Botanique

de Buitenzorg (1863) pi. 1.— Motley, J., Extract from a letter. Hook. J. Bot. & Kew Gard. Misc. 9

(1857) 148-153. — Smith, J. J., Trop. Natuur 18(1929) 156. — Solms-Laubach, H., Abh. Naturf. Ges.

Halle 13, 3 (1875) 40 pp., t. 24-27; Ann. Jard. Bot. Buitenzorg 9 (1891) 184- 246. — Teijsmann,

J.E., Nat. Tijds. Ned. Indie 1 (1850) 431-440; ibid. 12 (1856a)279-281; Hook. J. Bot. & Kew Gard.

Misc. 8 (1856b) 371-374; Ann. Hortic. Bot., Fl. Jard. Roy. Pays-Bas (1858) 27-30. — Teijsmann,J.E. & S. Binnendijk, Nat. Tijds. Ned. Indie 1 (1850) 425-430; Cat. Plant. Hort. Bot. Bogor. (1866)

15.— Vriese, W.H. de, Memoire sur R. rochussenii et patma (1853) 9 pp., 2 pi.

KEY TO THE SPECIES AND VARIETIES

1 a. Ramenta short, more or less pustulate, only a few millimetres long, in some cases

clavate 2

b. Ramentamore than 2.5 mm long 4

2a. Flowers 15-20 cm in diam. Opening of diaphragm (orifice) relatively wide, leaving

the lower inner side of the flower tube exposed to view from above. Anthers c. 15

7. R. manillana

b. Flowers at least 25 cm in diam. Orifice of diaphragm too small to expose the inner

side of the flower tube. Anthers 20-40 3

3a. Flowers (15-)30-60 cm in diam.Rim of disk steeply raised. Anthers (25-)32-40.

Ramenta very short knobs 9. R. patma

b. Flowers 26-30 cm diam. Disk rim only slightly raised. Anthers about 20

See R. borneensis, under Incompletely known species, p. 34

4a. Perigone lobes and diaphragm in general red or orange throughout, including the

warts. Diameterof flowers on average 20 cm (15-30). Ramenta slender and swol-

len at apex 5

b. Perigone lobes with white warty blots, or disk with many processes, or flowers

larger than 30 cm diam. (R. micropylora) 6

5a. Disk of flowers totally smooth or with up to 8 processes, rim not raised. Anthers

15-20 11. R. rochussenii

b. Disk with c. 25 processes, rim raised. Anthers20 13. R. tengku-adlinii


6a. Orifice of diaphragm angular or slightly lobed, very small compared to the size of

the flower, 3-9 cm in diam., too small to be able to count the c. 15 processes. Ra-

menta often branched and apices of branches globular, swollen. Flowers 30-60

cm diam.Warts on perigone lobes reddish 8. R. micropylora

b. Diaphragm opening relatively larger 7

7a. Apices of the ramenta either more or less distinctly swollen or crateriform...

8

b. Apices ofthe ramenta not distinctly swollen 10

8a. White warts across the base of the perigone lobes very large, 4 or 5 only. Ramenta

slender, apices swollen. Anthers c. 20. Diaphragm with only one basal ring of

dark-brown warts 4. R. hasseltii

b. White warts across the base of the perigone lobes more than 5. Diaphragm with

several concentric rings of warts 9

9a. White warts across the base of the perigone lobes 10-12. Some ramenta with

swollen, others with crateriform apices. Anthers c. 30 3. R. gadutensis

b. White warts across the base of the perigone lobes 7-9. Ramenta slender, all with

swollen apices. Anthers 20-25 2. R. cantleyi

10a. White warts in tangential direction about halfway along the perigone lobes rela-

tively covering less space than the red parts, c. 8 in number

See R. tuan-mudae, underIncompletely known species (p. 36)

b. White warts on perigone lobes more numerous 11

11a. Flowers c. 70 cm in diam. White warts on the perigone lobes very small, cover-

ing far less space than the red-colouredpart. Anthers (20?-)24-30

6. R. kerrii

b. White warts on the perigone lobes covering more space than the red-coloured

part 12

12a. White warts across the base of the perigone lobes 5 or 6(—10) 13

b. White warts across the base of the perigone lobes more than 10 14

13a. Flowers up to 80 cm diam. Lower face of the diaphragm with ramenta. Anthers

26-38 12. R. schadenbergiana

b. Flowers (16-)25-37.5 cm diam.Lower face of diaphragm without ramenta. An-

thers c. 20 10. R. pricei

14a. Larger whiteblots on perigone lobes interspaced with numerous small ones. Cir-

cular blots on diaphragm contrasting, isolated and with broad dark-brown margin

5. R. keithii

b. Larger white blots on perigone lobes not or very sparsely interspaced with smaller

ones. Circular whiteblots on the diaphragm little contrasting with their surround-

ing tissues, often more or less connected in a network of dark reddish bands and

with only a thin dark redbrown margin 15

15a. Ramenta covering the inner side of the flower tube from the rim ofthe diaphragm

downwards to the outer annulus la. R. arnoldii var. arnoldii

b. Ramenta missing in a zone above the outer annulus at the base ofthe column...

lb. R. arnoldii var. atjehensis


1. Rafflesia arnoldii R.Br.

Rafflesia arnoldiiR.Br., Trans. Linn. Soc. 13 (1821) 201; 19 (1845) 221, t. 22-26; Solms-Laubach,

Ann. Jard. Bot. Buitenzorg 9 (1891) 237, t. 27, 28; Koord., Bot. Overz. Raffles. Ned.-Indie (1918)

31, pi. 1, 2, 3; Coomans de Ruiter, Trop. Natuur 22 (1933) 165; 188 (as R. tuan-mudae); Meijer,

Nat. Geogr. 168 (1984) 136.— Type: Raffles & Arnold s.n. (K).

a. var. arnoldii

Mature buds up to 30 cm diam. Flowers (55-)70-100 cm in diam. Perigone lobes

with up to c. 15 whitewarts in radial and lateral direction, only occasionally interspersed

with smallerwhitewarts, the whitewarts occupying slightly more space than the reddish

brown background. Processes on the disk (20-)35-40(-50). Diaphragm with an open-

ing about 2/3 of the total diaphragm diameter, raised rather high at theend of flowering

phase, upper face light reddish pink with c. 40 radial series of 4 or 5 white circular warts

which are only a few mm in diameter, lower face just below the rim with 3 or 4 series

of whiterounded blots. Ramenta up to 10 mm long when mature, all over the inner side

of the flower tube, simple or forked and when fresh with papillose fluffy apices, never

crateriform swollen like in R. gadutensis. Maleflowers with 36-40 anthers.

Distribution — Malesia: Sumatra(Aceh, W Sumatra, Bengkulu, Lampong), Borneo

(W Sarawak, W Kalimantan).

Habitat — Primary and disturbed lower montane forests, up to c. 1000m altitude.

Ecology — Host plant most commonly Tetrastigma leucostaphylum (Dennst.) Alston

ex Mabb. [= T. lanceolarium (Roxb.) Planch.].

Notes — 1. See also underIncompletely known species, Rafflesia tuan-mudaeBecc.,

p. 36.

2. A plant of this species flowered on 9 February 1857 in the Bogor BotanicalGar-

den. See paragraph on ex situ cultivation, p. 20.

b. var. atjehensis (Koord.) Meijer

Rafflesia arnoldii R.Br. var. atjehensis (Koord.) Meijer, comb, et stat. nov. — Rafflesia atjehensis

Koord., Bot. Overz. Raffles. Ned.-Indie, Nieuwe Addenda (1918) 11; Bull. Jard. Bot. Buitenzorg 3

(1918) 77, 3 pi. — Type: Koorders 44060 (BO).

This variety differs only in the almost glabrous, 2 cm wide zone at the base of the

perigone tube, with only scattered, very short (3-6 mm long) ramenta.

Distribution— So far only known from the type locality Aceh, Locop. Possibly also

near Bohorok near the boundary of the Leuser National Park.

Habitat— LowlandDipterocarp forest.

2. Rafflesia cantleyi Solms-Laubach

Rafflesia cantleyi Solms-Laubach, Ann. Jard. Bot. Buitenzorg, Suppl. 3 (1910) 1. —Type: Cantley

s.n., received at Kew in 1887, fragments given to Solms in 1901,seen by the author in STR.

Rafflesia hasseltii auct. non Suringar: Meijer, Mai. Naturalist 36 (1983) 21, 2 col. photos, map; Wong,

Nature Malays. 15 (1990) 56-59, col. photos.


Flowers c. 30-55 cm across, in all respects very much like R. hasseltii but perigone

lobes with about 6-8 big whitish warts in radial and lateral direction and up to 10 in the

basal rows, sometimes these blots more or less confluent in bizarre wavy edged config-

urations. Diaphragm opening rounded or angular, almost half as wide as the total diam-

eter, 4-8 cm broad upper face light pinkish or whitish with about25 radial series of 3 or

4 dark red circled dots; lower face above the insertion with 3 concentric series of stalked

toadstool-like ramenta, with 5 mm long stalks and capped by 5-8 mm wide disks, to-

wards the rim 5 concentric series of oval white blots, 12-15 mm wide and 8-10 mm in

radial direction. Ramenta covering the inner face of the flower tube 10-12 mm long,

partly branched, with swollen apices, papillose grainy, lower down shorter, c. 2 mm

long. Maleflowers with 20-25 anthers.

Distribution—

Malesia: Malaya (Perak, Kelantan, Pahang, TiomanI., Kedah), see

map in Meijer, I.e. 1983, updated (in ms) by Matthew Wong. See also Meijer & Wong,

Mai. Naturalist 47 (1993) 10-11.

Habitat—

Lowland Dipterocarp forest and secondary forest.

Notes—

1. This species was up to 1984 consideredby the author to be identical with

R. hasseltii Suringar, following identificationby Ridley and others.

2. See also note 2 under R. hasseltii.

3. Rafflesia gadutensis Meijer

Rafflesia gadutensis Meijer, Blumea 30 (1984)211; Weevers in Koningsberger (ed.), Leerb. Alg. Plantk.

2 (1942) f. 186 (as R. arnoldii): Richards, Trop. Rain Forest, ed. 2 (1996) 155: f. 6.23; Meijer, Pal-

mengarten 60/2 (1997, '1996') 38-41. —Type: Meijer 17003 (BO holo; MO), W Sumatra.

Rafflesia spec.: Schaefer, Trop. Natuur (1940) 21-23;

Flowers 40-46cm diam. Perigone lobes with 10-12 whitish-pinkish blots along the

median and along the base, largest blots 2 by 1 cm, few small blots in between, some of

the larger sometimes merged and up to 5 cm broad. Diaphragm opening about2/3 of the

total width, pinkish red with about30 radial rows of circular light pinkish red margined

warts, lower face of diaphragm with 5 concentric rings of white blots close to the rim,

the two rings of warts closest to the perigone tube with flat or somewhat crateriform

oval fringed apex, c. 9 by 8 mm across, the tops supported by l-2(-6) mm long stalks,

making themlook like minusculetoadstools. Perigone tube c. 8 cm deep, covered at the

inside with rather fugaceous ramenta which are 6-8(-9) mm long and have swollen

heads 1 mm long and 2-3 mm wide, sometimes crateriform at apex, stalks c. 1 mm

across, a few branched or lobed ones near the diaphragm. Between the ramenta on the

inner side of the perigone tube some scattered toadstool-like protuberances close to the

inside of the diaphragm, but with only 1-2 mm long stalks. Maleflowers with c. 30 an-

thers.— Plate 1.

Distribution —Malesia: W Sumatra (Westcoast, Ophir District, Padang-Bukittinggi;

Bengkulu, Kayu Tanam, Padang-Solok, Ulu Gadut).

Habitat— Primary and secondary lower montane forest, around 350-600 m altitude.


Notes—

1. First seen by Korthals around 1834 and illustrated for him but unpub-

lished; also copies made for Solms-Laubach. Also found near the gold and silver mine

Simau, Bengkulu, and photographed there around 1934 by K. Schaefer [see Schaefer,

Trop. Natuur 29 (1940) 21-23].

2. Beccari collected the species in August 1878 near Ayer Mancior, the waterfall in

the Anei Canyon between Padang and Bukit Tinggi, at about 360 m altitude near Kayu

Tanam, West Sumatra. He and Solms-Laubach misidentifiedthis species as Rafflesia

arnoldii.

4. Rafflesia hasseltii Suringar

Rafflesia hasseltii Suringar, Acta Soc. Reg. Sc. Neerl. 25 Oct. 1879 (1880?) [4-5]; Assoc. Avanc. Sci-

ences, Congr. Alger (1881)621-626, pi. 11, 12; in Veth, Midden-Sumatra (1884) 13, t. 1,2; Solms-

Laubach, Ann. Jard. Bot. Buitenzorg 9 (1891) 23; in Engl., Pflanzenr. 5 (1901) 9, f. 1; Ann. Jard.

Bot. Buitenzorg 3, Suppl. pt 1 (1910) 1-4, pi. 1-6;Koord., Bot. Overz. Raffles. Ned.-Indie (1918)

76-81, pi. 12; Theunissen, Trop. Natuur 13 (1924) 150-152, 2 f.; Harms in Engl. & Prantl, Nat.

Pflanzenfam., ed. 2, 16b (1935) 264; Meijer, Grasduinen 5 (1991) 65, photo. — Type: Suringar (L).

Flowers c. 38-50 cm diam.Perigone lobes c. 11.5-13 cm long and 15-17 cm wide,

with about 5 large pustules across, these ranging from 5 by 3 to 10 by 1 cm. Perigone

tube c. 18-20cm wide, neck of column 3 cm. Processes 15-24, coloured like the disk,

light yellowish, but dark brown at the apex and not flattened as in some other species.

Diaphragm opening abouthalf as wide as apex of flower tube, pale whitish or yellowish

with a dark brown zone near the rim and a basal ring of rounded or oblong darkbrown

warts. Upper zone of the flower tube near attachment of the diaphragm with 4 rings

of toadstool-like compound ramenta, grading into white blots at the lower face of the

diaphragm. Numerous bristles on lower face of the corona (= crest of the disk) and all

along the column. Ramenta linear, with swollen apices. Maleflowers with 20 anthers.

Distribution — Malesia: Sumatra (W Sumatra, Riau, Jambi), Peninsular Malaysia.

Habitat— Primary and secondary lower montane forest, around 400-600m altitude.

Notes — 1. Of the six localities known four were located during the last 10 years.

2. This species is closely related to Rafflesia cantleyi and seems to hybridize with it

in the Malay Peninsula.

3. The description was composed from the original one and from a few fragments of

the type in L and some photographs. Most records come from pictures.

5. Rafflesia keithii Meijer

Rafflesia keithii Meijer, Blumea 30 (1984) 211; Salleh & Latiff, Blumea 34 (1989) 113.— Type: Ma-

dani s. n. (SAN holo, in spirit), Sabah, Sungai Melaut.

Rafflesia spec.: Corner in Luping et al., Kinabalu, summit of Borneo. Sabah Soc. Monogr. (1978) 136,

168, t. 154, photos.

Maturebuds up to 25 cm diam.Flowers on average c. 80(-94) cm in diameter (few

records). Perigone lobes with rather dense numerous whitewarts, small ones interspers-


ed with larger ones, the latter about 10-12 across the greatest breadth ofthe lobes. Dia-

phragm with5 concentric rings of white warts in about 40 radialrows, each surrounded

by a dark red brown margin, lower face of diaphragm with 5 or 6 rings of white blots,

those of the ring closest to the opening for a large part confluent, towards the transition

into the flower tube not with ramentae formed like white-headedtoadstools (stalked

disks), but uniformly with the same type of long ramentae as those on the inner side of

the flower tube. Perigone tube densely covered withramenta inside, the ramenta 5-6

mm long near the diaphragm and often fascicled (in bundles), only a few branched. Disk

11-13.5 cm in diam. Processes 38-42(-46), flattened.Annulus interior in a budof 22

cm diam., 2 mm wide, annulus exterior flat, 5 mm wide and grooved. Male flowers

with about 40 anthers and with densebristles on the lower face of the crest of the disk

and around the anther cavities. Femaleflowers with a swollen ring along the lower rim

of the crest of the disk, c. 3 mm high and broad. — Plate 2.

Distribution—

Malesia: Borneo (Sabah, most likely also in E Kalimantan, see R.

witkampii under Incompletely known species, p. 36).

Habitat—Upper Dipterocarp forest with Parashorea malaanonan,around500-600m

altitude.

Ecology — Host plant Tetrastigma leucostaphylum (Dennst.) Alston ex Mabb., very

possibly also T. diepenhorstii Miq. (= T. trifoliolatum Merr.) but proof only from the

former with a host plant with very low branches.

Notes — 1. At first sight this taxon comes close to R. arnoldii as defined in this revi-

sion. However, close study of the ramenta reveals that the single, toadstool-shaped ones

that are typical for arnoldii are missing in keithii (and in pricei, possibly the nearest rela-

tive of keithii). This view is supported by the occurrence of a hybrid at the Mamutcop-

per mine Rafflesia sanctuary in Sabah (Jamili Nais, oral comm.).

2. A main problem is still the lack of good documentationand sharp colourphoto-

graphs of Rafflesia specimens with mature flowers from E Kalimantan.

6. Rafflesia kerrii Meijer

Rafflesia kerrii Meijer, Blumea 30 (1984) 212; Meijer & Elliott,Nat. Hist. Bull. Siam Soc. 38 (1990)

117-133.— Type: Kerr 16980 (K holo;BM), Phuket.

Flowers up to 70 cm diam. Perigone lobes up to c. 20 cm long and 24 cm wide, with

scattered rather numerous warts, with 3-4 mm space between them, the warts smallest

relative to the red background ofany species known. Perigone tube with ramenta which

are mostly unbranched and only slightly swollen at apex, those near the diaphragm c. 10

mm long, those near the base of the tube 5 mm. Diaphragm only slightly lighter red than

the perigone lobes, about 30 cm in diameterwith a central opening about 20 cm across;

upper face with about 3 to 4 concentric rings of circular whitish, dark red ringed spots,

in about 30 radial lines; lower face with 7 concentric rings of white blots, which are up

to 7 mm across. Central part of disk with 32-40 flattened processes, 2-2.5 cm long.

Maleflowers with 25-30 anthers.


Distribution — Peninsular Thailand; Malesia: PeninsularMalaysia (Kelantan: Bukit

Tepoh; Kelantan-Perak border: Mt Chamag; Perak: Bintang Range).

Habitat— Lowland and hillDipterocarp forests, 500-1000 m altitude.

7. Rafflesia manillana Teschemacher

Rafflesia manillana Teschemacher, Boston J. Nat. Hist. 4 (1841) 63, t. 6; Ann. Mag. Nat. Hist. 9

(1842) 381; Solms-Laubach, Ann. Jard. Bot. Buitenzorg 9 (1891) 241, t. 26, f. 7-10; W.H.Br.,

Philipp. J. Sc, Bot. 7 (1912) 209, t. 12, 21; Merr., Spec. Blanc. (1918) 135; Enum. Philipp. Flow.

PI. 2 (1923) 120; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 16b (1935) 264; Pancho, Vase.

Fl. Mt Makiling (1983) 312, f. 98. — Type: Navarro specimen from Leyte (A, lost).

Rafflesia philippensis Blanco, Fl. Filip., ed. 2 (1845) 565.— Type: Navarro specimen from Laguna.

Luzon, now lost.

Rafflesia lagascae Blanco, Fl. Filip., ed. 3, 3 (1879) 231; most likely a nomen nudum.

Rafflesia cumingii R.Br., Trans. Linn. Soc. 19 (1845) 243.— Type: Cuming collection (BM), most

likely from the same site on Leyte as the Navarro collection seen by Teschemacher.

Flower buds up to 8 cm across, cupule 2.5-3 cm high, 4-6 cm wide, bud scales up

to 5.5-7 cm long. Flowers 15-20 cm wide and 7-9 cm high when expanded. Perigone

lobes light reddish brown, 4-6 cm long and 5-7 cm wide, white warts a mixture of

larger circular with oval ones, about 10-12 along a transverse line halfway the lobes,

mixed with numerous white dots. Diaphragm c. 1-1.4 cm broad, c. 7 cm in diameter,

strongly curved inwards along the opening, which is about 5/6th as wide as the apex

of the perigone tube; upper face rather densely randomly covered with oblongish white

warts. Disk of column c. 2 cm above the base of the perianth tube, 3.2-4.5 cm in diam-

eter including the rim, exposed while the flower is opened; rim of disk with a 3-5 mm

high raised part. Processes about 14-20(-30), sometimes almost missing in the centre

of the disk, 3.5-5.5 mm long, apex cylindric, with bushy cilia. Neck of column 1.3-

1.8 cm diameter, narrower in male flowers. One prominent annulus, 3 mm broad, 30-

38 mm diameteraround the base of the column. Perigone tube c. 2.5 cm deep, 5 cm

wide; ramenta at the inner side in about 12 rows, little stalked outgrowths, 0.5-1 mm

high, multilobedat apex, virtually absent near the base of the perianth tube, close to the

annulus; in a 1 cm wide zone below the diaphragm with short, warty lobed, whitish, 2-3

mm wide warts. Maleflowers with 10-15 anthers, relatively large for such small flow-

ers, not hidden in a sulcus as in the larger-flowered species, but attached rather close to

the lower margin of the disk; corresponding with each anther in the central column a

deep groove which runs down below the anther to the column base. Female flowers

with ovary 2.5 cm wide at apex, 1.0 cm high, above the annulus without grooves or

with very faint ones, less room under the corona (overhang) of the disk than in male

flowers, on its lower side a papillose stigmatic zone.

Distribution — Malesia: Philippines (Luzon, Samar, Leyte). Most of these records

are old history. The species is now rare and endangered.

Habitat— Lowland and hill forests, up to about 1000 m altitude.

Note — This is the smallest known Rafflesia, with flowers only about 15-20 cm in

diameter, a very narrow diaphragm, the whitish lobed ramenta only up to 1 mm tall and

male flowers only with 10-15 anthers.

Flora Malesiana, Ser. I, Vol. 13 (1997)28/1_2

Meijer. Sabah, Sabah, Mt Kinabalu. Photo Jamili Nais, 1990, donated 1994.Rafflesia keithiiPlate 2.

Meijer. Sumatra, Ulu Gadut near Padang. Photo W. Meijer, 1983.Rafflesia gadutensisPlate 1.

Meijer Rafflesiaceae 29/3_4

Meijer. Sabah, Tambunan. Photo Julius Kulip, 1989, donated 1992.Rafflesia priceiPlate 4.

Meijer. Sumatra, Mt Leuser National Park. Photo A. Rijksen.Rafflesia micropyloraPlate 3.


8. Rafflesia micropylora Meijer

Rafflesia micropylora Meijer, Blumea 30 (1984) 213.— Type: Koorders 44211 (Badings s.n.) (BO

holo),Lokop.

Rafflesia cf. atjehense auct. nonKoorders: W.J. de Wilde, WOTRO-Jaarboek (1972) 38, fig.

Rafflesia spec. 3 and spec. 4: Koord., Bot. Overz. Raffles. Ned.-Indie (1918) 109, 111. See note 2.

Flowers 30-60 cm diameter, no data availableon size of buds. Perigone lobes dark

orange-red all over, 16-18 cm long, warts light pinkish, little contrasting with their sur-

roundings, about 10 across at the base, with many smaller warts inbetween. Diaphragm

darker red than the perigone lobes, c. 20-25 cm diameter, upper surface with 3-5 con-

centric circles of circular reddish warts with darker red margins, weakly contrasting, in

about 40 radial lines; lower side of diaphragm with about 3(-5?) concentric circles of

flat circular knobs, c. 10 mm in diameter, on 5 mm high stalks, grading into the ramenta

towards the upperrim of the perigone tube; opening small, c. 3-9 cm wide, sexangular

or heptangular, sometimes weakly lobed. Column 5-6 cm high, c. 7-8 cm in diameter

at the neck. Disk 13-14cm diameter, with about 15 processes. Ramenta branched, with

swollen apices, c. 12 mm long, those towards the base ofthe perigone tube only 5-7 mm

long and unbranched. Maleflowers with 40 anthers.— Fig. 6; Plate 3.

Distribution — Malesia: Sumatra(Aceh). Protected inside the Leuser National Park.

Habitat — In lower montane forests around 300-500 m altitude.

Ecology — Host plant Tetrastigma leucostaphylum (Dennst.) Alston ex Mabb., and

possibly other species.

Notes — 1. The medium- to large-sized flowers, the small opening of the diaphragm

and the richly branched ramenta with swollen apices, make this species easy to recog-

nize.

2. The red-coloured diaphragm, together with the size and the small hole as illustrated

in Koorders (I.e.: plate 15, fig. G), make identificationof the two numbered species now

possible.

9. Rafflesia patma Blume

Rafflesia patma Blume, Flora 8 (1825) 609; Flora Javae 1 (1828)8, t. 1-3; De Vriese, Mem. sur Raf-

flesia Rochussenii et Patma (1853); Illustr. Rafflesia (1854) t. 5, 6; Rodigas, Fl. Serres 15 (1862-

65) 13, t. 1505-1508;Nagelvoort, Nat. Tijds. Ned. Indie 35, VII, 5 (1875) 171-180; Haak, Weekbl.

Pharm. 3 (1885) 19 pp., 2 pi.; Observ. Rafflesia patma (1889) 4 pi.; Solms-Laubach, Ann. Jard.

Bot. Buitenzorg 9 (1891) 238-239, pi. 16, f. 11-16; in Engl., Pflanzenr. 5 (1901)9, f. 2,5; Koord.,

Exk. Fl. Java 2 (1912) 179; Warb., Pflanzenwelt 1 (1913) t. 31, f. Dl-5; Hochr., Le Globe 57

(1918) 27-36, photo; Koord., Bot. Overz. Raffles. Ned.-Indie (1918) 50-61, pi. 6; Harms in Engl.& Prantl, Nat. Pflanzenfam., ed. 2, 16b (1935) 263; Backer & Bakh.f., Fl. Java 1 (1963) 165.

—

Type: Blume (L), Nusa Kambangan.

Rafflesia zollingeriana Koord., Bot. Overz. Raffles. Ned.-Indie (1918)67, pi. 10, 11; Backer & Bakh. f.,

Fl. Java 1 (1963) 165; Ervizal, Masters thesis (unpubl.) Agric. College Bogor (1989). — Type:

Koorders 40312 (BO), Java, Puger.

Flower buds prior to expansion up to 21 cm in diameter.Open flowers (30-)37-52

(-60) cm in diameter(in flowers described as R. zollingeriana sizes of 20-25 cm re-


ported). Perigone lobes flesh-coloured, brown-ocre or carmine red, c. 13—14(—19) cm

long and 10-14 cm wide, on the upper face with numerous whitish warts, up to about

25 at the largest breadth, surrounded by much smaller white warts, then same pattern as

in R. keithii. Diaphragm a 5-7 cm wide ring; opening in the centre 5-9 cm across; up-

per surface strongly pitted, glaucous or with clear circular whitish blots; lower face with

2 series of white oblong-angular blots, two to three times as long tangentially than radi-

ally, covered with ramenta or in some cases the region (2-)3-3.5 cm from the margin

of the opening and the zone close to the perianth tube devoid of ramenta. Disk carmine

red, upper face 6-6.5 cm from the bottom of the cupule, (7—)9—13.5 cm in diameter,

central part with (24-)37-44(-54) processes 2-3 cm long and 5-8 mm at base, coni-

cal or somewhat flattened; corona of the column-disk about 2-2.5 cm high and 1.5 cm

wide in radial direction, rim in some cases crenulate, papillose and fimbriate, directed

sideways under an angle of about 45°. Ramenta reduced to very short (up to 1 mm

long) tubercles, some branched. Maleflowers with about 25-32(-38 in S Sumatra)

anthers, 4-6 mm long and across, attached at the inner margin of the corona in a more

or less horizontal position, anther cavities 10-15 mm long and 8 mm broad, weakly

or not divided by two ridges, partitions between the anther holes provided with cilia.

Femaleflowers with a 2-4 mm broad, slightly papillose stigmatic zone divided by a

groove; rudimentary anthers c. 1—1.5 mm, in very shallow, broadly triangular cavities.

Gully at the neck ofthe column (sometimes referred to as Sulcus coronalis disci) about

8-9.5 cm in diameter, c. 17 mm from the rim of the corona. Annulus interior 4-6 mm

broad, well defined and sharply elevated (2-3 mm high), or only slightly raised, about

8-9.5 cm in diameter. Sulcus inter annuli 3-5 mm wide. Annulus exterior 3-10 mm

wide average and c. 1 mm thick, rather weakly developed, wrinkled like a rolled-up

blanket which is curved into a circle. Fruit with cupule 11 cm in diameter, 7-8 cm high,

diameter in upper part 7.5 cm, ovarial part 1-2.5 cm high and 5-10 cm wide, side of

column with about 40 grooves. Ripe seeds 0.8-0.9 mm long, 0.4 mm thick, light-

brown to dark-brown.

Distribution— Malesia: S Sumatra(Lampong, now extinct?; C and E Java (close to

extinction), possibly formerly also in Bali.

Habitat— Most localitiesare along the south coast of Java in remnants of forest be-

low 400 m altitude.

Ecology — Host species most commonly Tetrastigma leucostaphylum (Dennst.) Al-

ston ex Mabb., but possibly also T. glabratum (Blume) Planch.

Notes — 1. A medium-sizedRafflesia, at first sight much like R. arnoldii but the

lower side of the diaphragm often covered with ramenta. Warts on the perigone lobes

are a mixture of larger and smaller ones, more or less as in R. keithii from Borneo. The

flower tube is more or less covered with very short knob-like ramenta, not more than

one mm long, in some forms partly missing.

2. A form from East Java, with smaller flowers and ramenta almost absent in the

lower part of the flower tube, was described by Koorders as R. zollingeriana. Ervizal

(1989) found there in 1989 still viablepopulations.


10. Rafflesia pricei Meijer

Rafflesia pricei Meijer, Blumea 30 (1984) 214; Beaman & J.H.Adam, Sabah Soc. J. 7 (1984) 208-212;

Beaman c.s., Amer. J. Bot. 75 (1988) 1148-1162; Mat Salleh, Rafflesia magnificent flower of

Sabah (1991), several pi.; Meijer, Grasduinen 5 (1991) 63. — Type: Price s.n. (K holo), Mt Kina-

balu.

Rafflesia schadenbergianaauct. non Goeppert: Meijer, Kosmos 5 (1982) 60, fig.

Rafflesia tuan-mudae auct. non Becc: Anderson, Bioscience 26, 10 (1976) cover plate; Weber, Sabah

Soc. J. 3 (1967) 11, t. 1, 2; Yong, Magnificent PI. (1981) 2, fig.; Bailes, Kew Mag. 2 (1985) 275,

fig.; Jacobs, Trop. Rainforest (1988) 241.

Rafflesia spec.: Dorst, Avant que nature meurt (1965) 153, t. 10; Price, Proc. Cotteswold Nat. Field

Club 35 (1968) 93; Morley, Wild Flowers of the World (1970) t. 117C; McKinnon, Borneo (1975)

22, fig.; Bellamy, Botanic Man (1978)60, fig.; Ayensu, Jungles (1980) 49; Masni, Bt Jubil, Kajian

Ekologi dan taksonomi Rafflesia di Sabah khususnya di Tambunan,Thesis Univ. Kebangsaan Ma-

laysia (1984); Beaman & J.H. Adam, Sabah Soc. J. 7 (1984) 208.

Mature buds c. 15-20 cm in diameter. Flowers (16-)25-30(-45?) cm across. Peri-

gone lobes 7-10 cm long and 6.5—14(—19) cm wide, with 10-15 warts at base, many

of those wider in the tangential than in the radial direction, some confluent. Perigone

tube c. 14 cm wide. Column 3-4 cm high, c. 4-5(-9.5?) cm in diameter at the neck.

Disk 8.5-9 cm diameter, with 20-40 flattenedprocesses which are about 1-3.5 cm

long and up to 1-1.5 cm in diameter.Diaphragm (7—)12—13(—17) cm diameter, opening

c. 4.5-8.5 cm in diameter, slightly angular and with a white rim surrounded by a dark

brown margin; upper face of diaphragm with four concentric rings of warts, those in the

third ring from the centre sometimes linear, in the second ring more circular and especi-

ally in these two rings surrounded by dark brown margins; lower face of diaphragm

with 4 or 5 concentric rings of laterally stretched whitebands, withoutramenta. Ramenta

near the diaphragm in fascicled groups, slender, only slightly widened at apex, 2-6 mm

long near the insertion of the perigone lobes and 6-7 mm at the base of the tube. Male

flowers with 20 anthers.—

Plate 4.

Distribution — Malesia: Borneo (Sabah, Sarawak, Brunei, also Kalimantan?). Appar-

ently most populations are on MtKinabalu and the Crocker Range and Schwaner Range

in Sarawak.

Habitat— Upper Dipterocarp forests with Shorea platyclados and lower montane

forests, about 400-1300m altitude.

Ecology —Host species Tetrastigma leucostaphylum (Dennst.) Alston ex Mabb.

Notes — 1. This species apparently is now also known from Sarawak (Serian area)

and it is possible that a recent discovery of Rafflesia by Danna J. Leaman (University

of Ottawa) in the Upper Kajan near Long Ampin (East Kalimantan) also belongs to this

species.

2. Probably the first collection was made by Joseph and Mary Clemens (nr 28512),

on 17 March 1932, near Dallas on MtKinabalu. However, we are not quite sure because

the specimen does not show the pattern of the warts on the perigone lobes and the size

of the flower is almost too large for this species, though the label records "diameter 13-

18 inches" (= 32.5-45 cm). Meijer (in Dorst 1965) was the first to publish a photograph

of this species on Mt Kinabalu.


11. Rafflesia rochussenii Teijsm. & Binn.

Rafflesia rochussenii Teijsm. & Binn., Nat. Tijds. Ned. Indie 1 (1850) 425-430, t. 1, 2; 2 (1851) 651-

655; De Vriese, Mdm. sur Rafflesia Rochussenii et Patma (1853) 4; Illustr. R. Rochussenii (1854)

t. 1-3; Solms-Laubach in Engl., Pflanzenr. 5 (1901) 10, f. 6, 7; Ann. Jard. Bot. Buitenzorg, Suppl.

2 (1898) 3-12, t. 1; Ernst & Schmid in Ann. Jard. Bot Buitenzorg II, 12 (1913) 1-58; Koord., Bot.

Overz. Raffles. Ned.-Indie (1918) 82, incl. var. subaculeata Koord., I.e. 87; Steenis, Trop. Natuur

30 (1941) 179-181. — Type of species and variety: from W Java, lost.

Matureflower buds just before opening c. 10-13 cm in diameter.Flowers (15-) 17-

20(-30) cm in diam., c. 12 cm high. Perigone lobes c. 6.5-8(-9) cm long and 8—9(—11)

cm wide, 7-10 cm at insertion. Diaphragm c. 11 cm in diam., 2.5-4 cm broad, lower

surface covered with short (1-2 mm long) ramenta. Disk c. 5.5-6 cm above the base of

the cupule, (6-)7-9 cm in diameter, ivory white when fresh, flat or slightly raised in

the centre, rim sharp, not raised, centre without processes or with only 1-8 ('var. sub-

aculeata'). Sulcus below the disk c. 5 cm in diam. Neck of column4.5-5.5 cm in diam.

Perigone tube 12 cm wide in the middle between annulus interior and diaphragm; annu-

lus interior6.5-7 cm in diam., rather sharp, 2-4 mm thick; annulus exterior a flat 6-8

mm broad zone without ramenta. Ramenta long stalked subdiscoid knobs, c. 10 mm

long near the base of the perigone tube. Maleflowers with 15-20 anthers. Femaleflow-

ers with stigmatic ring distinctly narrowed. Fruits c. 8.5 cm wide and 7 cm high, ovary

6 cm wide and 3 cm high, disk decayed, cupula (5-)7-8 cm diam., 1.5-2 cm high.

Distribution — Malesia: W Java, N Sumatra (Mt Leuser and further south in Tapa-

nuli). In 1989 and 1990 two sites were rediscovered on Mt Salak, Java, and one around

1995 on Mt Gede, Java (collected by Arindansyah, oral comm. by H. Wiriadinata). The

species probably still occurred in Berastagi, Sumatra, in 1980. The site was, however,

logged and destroyed in 1981.

Habitat —- Lower montane forests up to about 1500 m altitude.

Notes — 1. The most distinctive character is the disk of the column with the rim only

slightly raised and the surface bald or only with a few processes, never more than eight.

The ramenta are slenderand in general have knob-like apices.

2. Cultivated in the Leiden Botanical Garden in 1851 from dug-up plants (see para-

graph on ex situ cultivation, p. 20) and illustrated from damaged flowers by De Vriese

in 1853 and 1854.

12. Rafflesia schadenbergiana Göpp.

Rafflesia schadenbergianaGopp. in Hieron., Bull. Internat. Bot. Horticult. St. Petersb. ('1884', 1885)

35-36, 1 pi.; Gartenflora 34 (1885) 3-7, pi. 1177; Ueber Rafflesia schadenbergiana Goepp., Ein

Beitrag zur Kenntnis der Cytinaceen (1885) 1-10, t. 1, 2; Solms-Laubach, Bot. Zeitung (1885) 507;

Ann. Jard. Bot. Buitenzorg 9 (1891) 189.— Type: Hieronymus s.n. (K, WRSL iso?).

Flower buds prior to expansion 16-20 cm in diameter, cupule 10-14 cm in diameter,

bracts up to 17-18 cm long, 12-13 cm wide. Open flower about 80 cm diameter. Ra-

menta on the insideof theflower tube 7-10 mm long, filiform, somewhat thickened at

apex, or branched, partly in fascicles. Diaphragm 6-8 cm from insertion to opening,


opening 13-14cm in diameter, margin with pinkish zone, lower face except the marginal

zone provided withramenta c. 4-5 mm long. Perigone lobes 25-26 cm diameter, with

yellowish whitish warts, which are laterally stretched, irregular shaped, partly connected

and about 4 or 5 across in the middlepart. Disk 12-13 cm diameter, processes 30-50.

Maleflowers with 26-38(-40) anthers.

Distribution — Malesia: Philippines (Mt Apo, Mindanao). No records during the 20th

century.

13. Rafflesia tengku-adlinii Salleh & Latiff

Rafflesia tengku-adlinii Salleh & Latiff, Blumea 34 (1989) 111-116; Fl. Males. Bull. 11 (1995) 425-

428. — Type: Mat Salleh c.s. KMS 2180 (UKMS holo),Trus Madi Mts.

Matureflower buds about 13 cm in diameter.Flowers rather small, c. 20-24 cm in

diameter.Disk about 6 cm diameterand neck of column 3 cm. Perigone lobes and dia-

phragm orange throughout including randomly distributed roundish warts, except near

the opening, the diaphragm progressively dark reddish towards the centre. Diaphragm

about 12.5 cm wide, with a 5 cm wide aperture. Ramenta slender, 3-5 mm long, partly

branched and always with swollen apices, density somewhatreduced towards the open-

ing and at the base of the tube, also present on the lower face of the diaphragm. Disk c.

7 cm diameter, flattenedprocesses 25. Maleflowers with 20 anthers.

Distribution — Malesia: Sabah, eastern slopes of Mt Trusmadi. Locality destroyed

by loggers a few weeks after the plants were discovered. Lateralso found in the Gunung

Lutong area at 600 m altitude inside the Yajasan Sabah concession.

Most likely the same species as the one seen by Meijer around 1964 near Telupid as a

decaying flower.

Note — The measurements of the column, neck and disk were inferred from the

drawing with the original description.

INCOMPLETELY KNOWN SPECIES

Rafflesia borneensis Koord., Bot. Overz. Raffles. Ned.-Indie (1918) 47, pi.4. —

Type: Witkamp collection(BO).

Relatively smallflowers up to only 20-30 cm in diameter.Flower tube very flat and

disk of column with a rather flat rim. Ramenta as far as seen in the specimen very short

and sparse; Koorders described them as very short, rarely longer than 1 mm, with nar-

row pointed apex. He was unable to describe the perigone lobes and the diaphragm.

Distribution— Borneo, only once found in NE Kalimantan, Sekerat Mts, in 1917.

Habitat— At 600-700m altitude, according to Coomans de Ruiter, Trop. Natuur22

(1933) 166.

Note—

Because of the flattened rim of the disk this species is rather similar to R.

rochussenii. It is, however, distinguishable by the possession of the processes on the

disk and by the very rudimentary ramenta, like those of R. patma. The remaining mate-

rial in alcohol is very fragmentary and a good complete description is not possible.


Rafflesia ciliata Koord., Bot. Overz. Raffles. Ned.-Indie (1918) 64, pi. 9. — Type:

Witkamp collection from Sekerat Mts, NE Kalimantan, no materialfound in BO.

All we can learn about this enigmatic species from the original description, based on a

bud of 17 cm diam., is that the perigone lobes have numerous dense warts, the ramenta

are simple or branched, not swollen at apex, the lower face of the diaphragm has only

bristles, no warts, and the disk has many processes, pointed at the apex. The hairy lami-

nae between the anther cavities and the shape ofthe anthers and manyother detailsgiven

by Koorders are shared by most species ofRafflesia and are ofno diagnostic importance.

A collection from the same huge general region of the Sekerat Mts was madeon the

Sangkulirang Peninsula by Kostermans, Sept. 1957, near Mapula(?) vilage. It consists

of a male bud of 14 cm diameter with 40 anthers. This would make it likely that this

specimen either belongs to R. keithii or that R. arnoldiioccurs all the way from SW and

C to NE Kalimantan. In case future fieldwork on the Sekerat Mts would show that R.

keithii is common therewe have enough circumstantial evidence that specimens of that

species were described by Koorders under the names R. witkampii and R. ciliata. The

latter species name could be excluded from priority because of the lack of a convincing

type specimen. It is not possible to solve this puzzle without the collectionof new evi-

dence in the field.

Rafflesia titan Jack, Appendix Descr. Mai. Plants I (1821), reprinted in Hook. Comp.

Bot. Mag. 1 (1835) 259.

The reason William Jack made his own description of Rafflesia and gave the name

R. titan to the plants collected for Raffles and by Raffles and him in the province of

Bengkulu before he died on 15 September 1822 was revealed in a letter he wrote in May

1821 to Wallich in Calcutta. The letter was reprinted by Burkill in a paper in J. Straits

Br. Roy. Asiat. Soc. 73 (1916) 148-268, and contains the following statement: "You

must observe that though labelled an appendix to the Malayan Miscellanies it has been

kept back till we hear what is done at home about the great flower (Rafflesia). If it is

brought forward in England then this is to be suppressed and not published; if no, then

this may be used in the event of the French getting hold of it, as a proof of priority of

publication."

From a letterhe sent home 10 April 1820 [published by W.J. Hooker in Hook. Comp.

Bot. Mag. 1 (1835) 135] we learn: "I lately sent to England a short account of some of

my most interesting plants, to be noticed there in some way or other, including the su-

matran gigantic flower, my two new Pitcher-Plants, the Camphor, the Sago and a new

genusof mine, which Sir S. Raffles has forwarded for me, with the drawings of them,

to Mr. Marsden, to make such use of themas he thinks best."

It must have taken sail ships at least three months to reach England from Sumatra.

On 30 June 1820 Robert Brown read his famous paper on Rafflesia at a meeting of

the Linnean Society of London and an excerpt of this was published November or De-

cemberof that year in the Dutch magazine Algemene Kunst en Letterbode vol. 2, nr. 3,

and probably also in the German journal Flora.

36 Flora Malesiana, Ser. I, Vol. 13 (1997)

From the correspondence between Dr. D.J. Mabberley, Dr John Bastin and myself in

1984and 1985 no convincing evidence has been produced that Jack's Appendix reach-

ed England before the Brown paper came from the press in May or June 1821 and was

distributed at least to various scientific quarters. The most likely date of publication of

Brown's paper should be June 1821 at the latest, not July, the earliest month of arrival

of two reprints in India [see Merrill, J. Arnold Arbor. 33 (1952) 203]. A copy sent to

Brown by Jack was mailedin a letter dated 23 May 1821 according to Mabberley in an

unpublished note, a part of the correspondence, in which he wrote me (4 March 1985):

"it would be a service to the literate world to have this name (R. arnoldii R. Brown)

fixed for all time."

To take any further doubt away we can also argue that we now know that before 1940

there were at least two species of Rafflesia known from Bengkulu (see under R. gadu-

tensis in this treatment) and that the description by Jack is not fully convincing us that he

had not mixed data from two species in his species description and that only for R. ar-

noldii we have very clear drawings and a type specimen available to proof its true identity.

Rafflesia tuan-mudaeBecc., Atti Soc. Ital. Sc. Nat. 11 (1868) 197; Nelle For. Borneo

(1902) 166, illus.; Solms-Laubach, Ann. Jard. Bot. Buitenzorg 9 (1891) 239, t.27,

f. 4, 5, 9; Koord., Bot. Overz. Raffles. Ned.-Indie (1918) 66; Pawozan, Family

Tree, For. Dept. Sarawak Internal Newsl. 18 (1988) 3-7, col. phot. — Type: Bec-

cari s. n. (FI-B).

Medium-sized flowers c. 44-56 cm diameter. Lower face of diaphragm and ramenta

much like R. arnoldii but perigone tubes with only about 5-8 whitish warts across the

broadest part, these relatively smaller than those of R. gadutensis. Number of anthers

not known.

Distribution — Malesia: W Sarawak (Mts Pueh, Gading, Rara).

Habitat — As described by Beccari (1902) in lowland Dipterocarp forest.

Notes — 1. Beccari and Solms-Laubach did not have a sharp concept of this entity.

Beccari himself [Nuovo Giorn. Bot. Ital. 7 (1875) 70-75] reduced his species to R.

arnoldii Brown but later on left it to Solms-Laubach (I.e. 1891) to revive it, mainly

based on the smaller size of the flowers and fewerblots across the perigone lobes.

2. Coomans de Ruiter collectedRafflesia in SW Borneo, later in Bogor identified as

R. tuan-mudae,but the flowers illustrated by him are good matches of R. arnoldii. How-

ever, the smaller form described by Beccari was lateralso foundon Mt Rara by Dr. J.

Mogea (BO) and Dr. W.J. J.O. deWilde (L). So it now appears to be a form restricted

to three isolated mountains. As a widespread species R. arnoldii has greater variability in

size and form than some species with smaller areas.

Rafflesia witkampii Koord., Bot. Overz. Raffles. Ned.-Indie (1918) 61, pi. 7, 8. —

Type: Witkamp collection, Sekerat Mts, NE Kalimantan (BO, in spirit).

This 'species' was described by Koorders merely on the basis of the study of a bud,

25 cm diameter, of a female flower. The column was expanded and a rupture in the tis-


sues between the annuli produced the 'wings' illustrated. Most likely the bud was left to

dry out and later returned to a bottlewith spirit. Several other 'distinctive' characters have

no value because they were described from an immature bud. For the timebeing it is

impossible to interpret this species as anything else than perhaps the female flowerof R.

ciliata Koord., collected at the same locality, an enigma itself. From the size of the bud

we can exclude R. pricei as a possible identification. Rafflesia keithii would fit the size.

The species is possibly still present at the type locality though the site is doomed to

disappear because of the planned establishmentof a cement factory at that place. Other

sites may still survive east ofthe Kutai National Park and in and around Sangkulirang.

Ultimately the name may deserve priority over R. keithii.

RHIZANTHES

Rhizanthes Dumort., Anal. Fam. PI. (1829) 14; Meijer & Veldkamp, Blumea 33 (1988) 329-342;

Meijer in Kubitzki (ed.), Fam. & Gen. Vase. PI. 2 (1993) 561. — Brugmansia Blume in Van Hall,

Bijdr. Natuurk. Wetensch. 2 (1827) 422, nom. illeg., non Pers. (1805). — Zippelia Rchb., Handb.

Nat. Pfl.-Syst. (1837) 164, nom. illeg., non Blume ex Schult. & Schult. f. (1830). — Mycetanthe

Rchb., Deut. Bot. Herb.-Buch (1841) 61, nom. illeg., superfl. — Type species: Rhizanthes zippelii

(Blume) Spach.

Flowers unisexual or bisexual, buds pyriform, from a basal cupule attached to or

arising from the roots or basal stem and aerialroots of the host, surrounded by three or

four whorls of 5 scales, these sessile, imbricate, concave, more or less ovate, entire,

brownish. Perigone lobes 16, valvate, with worm-like apices attached inside a cavity

(crater) of the central column. Anthers in a ring around the lower margin of the column

disk, about 38-50, each with 2 superimposed loculi. Male flowers with a rudimentary

ovary, mature perigone tubes campanulate, radially striped in the center. Fruits hidden

under dried-out flowers. Seeds with the same structure as those of Rafflesia [see Bou-

man & Meijer, PI. Syst. Evol. 193 (1994) 187-212]. — Fig. 10-12.

Distribution— In Malesia 2 species (Sumatra, Peninsular Malaysia, Java, Borneo).

Ecology — The host of both species is in general Tetrastigma papillosum (Blume)

Planch., sometimes also T. leucostaphylum (Dennst.) Alston ex Mabb. [= T. lanceo-

larium (Roxb.) Planch.].

Note — A very clever field investigation into the pollination ofRhizanthes (most like-

ly R. lowii) in S Thailand and the Malay Peninsula (Perak) was reported by Banziger

(1996). He observed 900 visits of Calliphoridae (Bow flies) to 270 flowers and consid-

ers 6 species to be pollinators and the others non-pollinating visitors. His conclusion is

that Rhizanthes just like Rafflesia mimics carrion in a pollination syndrome based on

brood-site deception. Apparently eggs can be laid but the larvae starve to death. Insects

are also lured to the nectaria at the base ofthe bayonets. As known to all observers, the

flowers ofRhizanthes smell to the human nose much fainter than those of Rafflesia. See

for other interesting details the original publication.

Reference: Banziger, H., Nat. Hist. Bull. Siam Soc. 44 (1996) 113-142.


KEY TO THE SPECIES

la. Flowers red-brown at anthesis, unisexual. Basal darker part of the bayonets cover-

ed with a velvety layer of c. 1.5 mm long, strongly branched antler hairs

l.R. lowii

b. Flowers white at anthesis, bisexual, malewith rudimentary ovary, or male. Basal part

of the bayonets with short hairs only once or twice branched or only curved at apex

2. R. zippelii

Fig. 10. Rhizanthes lowii (Becc.) Harms. a. Flowerbuds and open flower on stem or woody root of

The mature bud left shows the cupula at base, the bracts, and the 16 valvate perigone lobes.

The open flower shows the perigoneand the narrow bayonets which in the bud were directed downwards

and partly hidden in an apical cavity of the column; b. mature male flowerbud with rudimentary ovary;

c. column of male flower, showing ring of anthers; d. section through the former; e. female flower-

bud; f. column of female flower, stigmatic ring below the apex; g. ramenta from the glandular bases

of bayonets from where they are often removed by frequent fly visits. Drawings by Janis Atlee, from

photographs by W. Meijer, Kinabalu National Park, 1994.

Tetrastigma.


1. Rhizanthes lowii (Becc.) Harms

Rhizanthes lowii (Becc.) Harms in Fedde,Rep. 36 (1934) 287; Corner & Watanabe, Illus. Guide Trop. PI.

(1969) 163. — Brugmansia lowi Becc., Atti Soc. Ital. Sc. Nat. 11 (1868) 198 ('lowi '); Koord., Bot.

Overz. Raffles. Ned.-Indie (1918) 101, t. 17, f. C- E; Merr., Enum. Born. PI. (1921) 244; Ridl.,Fl.

Malay Penins. 3 (1924)20, t. 137.— Brugmansia zippelii Blume var. lowii (Becc.) Hook. f. in DC.,

Prodr. 17 (1873) 113.— Mycetanthe lowii (Becc.) Hochr., Candollea 4 (March 1930) 188; Steenis,

Bull. Jard. Bot. Buitenzorg 111, 11 (June 1930) 18. —Type: Low s.n. in Herb. Beccari (FI holo; B).

More extensive list of references in Meijer & Veldkamp, Blumea 33 (1988) 335.

Flowers always unisexual, concolorous white to reddish (on Mt Leuser also with a

white tube and brown lobes). Buds ovoid-pyriform, up to 12 by 7 cm; open flowers

infundibuliform, 18-35 cm in diam. Perigone lobes 6-12 mm wide at base, gradually

narrowed, basal hairy part c. 3 cm long, gradually narrowed into the geniculate, c. 2.5

cm by 1-2 mm apical appendage, the upper part of c. 6.5 cm without long hairs. Peri-

gonetube inside long hairy below, the upper c. 2 cm with short hairs with an antler-like

shape, male flowers with 50-60 lines, corresponding to the number of anthers. Central

column c. 2 cm long, in female buds with a c. 0.5 by 1 cm diameter stipe, and a globu-

lar more or less depressed apex, 1-1.5 by 1.5-2 cm diameter, apical cavity c. 1 cm

wide and deep. Male flowers with a c. 7 cm broad ring of 50-60 anthers around the

base of the globular disk; stipe with c. 50-60 ridges; ovary rudimentary. Columnar disk

of female flowers with a c. 7 mm wide stigmatic hairy zone along base; stipe not ridged;

ovary c. 1.5 by 3 cm diam. Fruit not seen. — Fig. 10, 11.

Fig. 11. Photograph W.J.J.O.

de Wilde, 1972.

Rhizanthes lowii (Becc.) Harms. N Sumatra, Ketambe (de Wilde 12148).


Distribution— Sumatra (Aceh, Gajolands, Sibolangit, Bengkulu, Palembang, Lam-

pong), Peninsular Malaysia (Pahang, Perak, Trengganu), Borneo (Sarawak, Sabah, W

and E Kalimantan).

Habitat— Lowland Dipterocarp and hill forest, 200-1500m altitude.

Ecology — Parasitic on Tetrastigma papillosum (Blume) Planch.

Notes — 1. Although in the footsteps of Beccari the epithet is often written as lowi;

ICBN Art. 73.10 and Rec. 73 C 1 make it clear that lowii is the correct form.

2. Banziger, Nat. Hist. Bull. Siam Soc. 43 (1995) 337-365 argues that there is only

one species (R. zippelii) in this genus. However, he most likely never saw the real

R. zippelii. Hebased his concept of it on the wrong identification by Meijer & Veldkamp

(1988) of a specimen collected by Molesworth in a locality in Malaya which Banziger

also visited himself.

2. Rhizanthes zippelii (Blume) Spach

Rhizanthes zippelii (Blume) Spach, Hist. Nat. Veg. 10 (1841) 554; Backer & Bakh. f., Fl. Java 1 (1963)

166. — Brugmansia zippelii Blume in Van Hall, Bijdr. Natuurk. Wet. 2 (1827) 422; Miq., Fl. Ind.

Bat. 1, 2 (1859) 684; Koord., Bot. Overz. Raffles. Ned.-Indie (1918) 94, t. 15, f. A-D. — Zippelia

brugmansia Rchb., Handb. Nat. Pfl.-Syst. (1837) 164, nom. illeg., superfl. — Mycetanthe zippelii

(Blume) Hochr., Candollea4 (March 1930) 188; Steenis, Bull. Jard. Bot. Buitenzorg III, 11 (June

1930) 18.— Type: Zippel s.n. (L holo, spirit).

Brugmansia bakhuizenii Heinr., Denkschr. Akad. Wiss. Wien, Math.-Naturw. KI. 78 (1906) 63, 66, t.

1, f. 1-4, t. 3, f. 3-5, 7, nom. prov., illeg.; Backer & Bakh. f., Fl. Java 1 (1963) 166, in obs. —

Voucher: Bakhuizen van den Brink s. n. (IB).

More extensive list ofreferences in Meijer & Veldkamp, Blumea 33 (1988) 339.

Flowers unisexual or bisexual, buds up to 10 by 5 cm, pinkish, smooth, at anthesis

first concolorous white, laterbright red or brown, 8-21 cm in diameterincl. the worm-

like apical appendages. Perigone lobes c. 2.5 by 1.5 cm at base, free part 4.5-5 cm long

(excl. the appendages), apex c. 0.5 cm thick; in upper c. 1.8 cm with short brown hairs

with antler-shaped apices; lobes curved downwards touching the substrate, initially

united at base in groups of 3 or 4, later free, splitting on the outside to c. 1.5 cm from

the base of the tube, on the inside to c. 3 cm, forming a thin c. 1.5 cm long membrane.

Perigone tube 2-2.5 cm long, inside with 46-55 brown lines around the column, fading

out towards the base of the lobes; base and insideof the tube with 5-7 mm long hairs,

apically slightly hooked; column c. 2 cm long, base narrow, c. 1 cm thick, head globu-

lar, 1.8-2 cm wide, apex purplish brown, hairy, crater 0.5-0.9 cm diam. Anthers 38-

50. Ovary with stigmatic ring c. 0.5 cm high, white. Fruit subglobose, up to 6.5 cm

diameter, brown, hidden by the dark-brown dried perianth. Seeds oblong, c. 0.75 by

0.3 mm, appendage oblong, about half as long as the seed. — Fig. 12.

Distribution —Sumatra (Aceh, W Sumatra, Bengkulu, Lampong), Peninsular Malay-

sia (Perak), Borneo (W Kalimantan, from Sabah and Sarawak no firm records), W Java

(Ciapus canyon, Mt Salak).

Habitat— Primary or secondary forests, often along streams on deep alluvial soils,

500-1500m altitude.


Ecology — Host is usually Tetrastigma papillosum (Blume) Planch., occasionally T.

leucostaphylum (Dennst.) Alston ex Mabb. (names based on herbarium study by the

present author). Van der Pijl [Trop. Natuur 22 (1933) 55] reported a species of Ville-

brunea sp. (Urticaceae)i as host.

Notes — 1. Blume's original illustration of the flower reproduced repeatedly in the

literature has given rise to some confusion because the flower shown had not yet fully

expanded, the perianth lobes still sticking together in groups of 3 to 5.

The first illustrations ofRhizanthes lowiishowed flowers with all perianth lobes free

from each other. Heinricher [Denkschr. Akad. Wien, Math.-Naturw. Kl. 78 (1906) 63,

66, illus.], who found R. zippelii in its natural situation, was considering to describe it

(Blume) Spach. a. Open flower, wine-red in colour; b. male flower in length

section, ovary rudimentary, no trace of stigma ring around the column; c. bisexual flower in length

section, ovary below the column which shows anthers as well as a stigmatic ring above them; d, e.

detail of the column, with stigmatic ring; f. ripe fruit, partial cross section, thousands of seeds attached

to the convoluted parietal placenta; g. ramenta at the base of the bayonets. Drawing by Janis Atlee:

d, e after Blume, g after Heinricher; others from photographs made by W. Meijer in Sumatra, Ulu Gadut

forest.

Fig. 12. Rhizanthes zippelii


as a new species (R. bakhuizenii ?). Probably Blume's flower was produced in the lab-

oratory after the host and buds were dug up from thejungle. There is nothing to indicate

that the flower from Mt Salak represented a differentspecies than those later discovered

at all other localities.

2. Backer & Bakhuizen f. (1963) distinguished two forms: R. zippelii s.s. with white

or flesh-coloured perianth, etc. from the Salak, and 'Brugmansia bakhuizenii’ with bright

red perianth, etc. from Mt Gede, Cidadap, and Garut. The latter, however, is nothing

else than the phase shown by flowers on the second or third day of flowering.

3. Since 1940 there have been no more reports of this species from Java (pers. obs.,

Kostermans pers. comm.). In October 1981 Meijer saw still some specimens of Tetra-

stigma papillosum in the steep Ciapus Canyon on Mt Salak. Cultivation and tree planta-

tions had replaced the original forest at the margin of the canyon. No botanists have

penetrated into the canyon in the past 60 years, but Meijer found in August 1996 that

guides are leading parties into it.

flora i,...flora malesiana,series i,volume 13 (1997)1-42 1 rafflesiaceae w.meijerlexington, u.s.a.)1...

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